Sea barley (Hordeum marinum ssp. marinum) is a wild Triticeae species with notable tolerance to salinity and waterlogging, yet its genetic resources remain poorly characterized. In this study, we assessed the transferability of 111 wheat and barley-derived simple sequence repeats (SSR) markers to H. marinum and evaluated genetic diversity and population structure across ten natural Tunisian populations (150 lines). A total of 38 primers (34.23

Water deficit is a major abiotic constraint limiting maize growth and productivity worldwide. Although silicon (Si) is not classified as an essential element, its beneficial effects on numerous crop species are well documented. Silicon has been shown to promote plant growth and enhance tolerance to abiotic stresses, particularly drought stress. Seed priming, a pre-sowing technique known to stimulate early germination processes, has emerged as a promising approach to enhance seedling establishment and stress tolerance in crops. In the present study, silicon-based seed priming was investigated as a strategy to alleviate the adverse effects of water deficit in maize (Zea mays) using two sodium silicate priming-solution concentrations (10 and 20 mM). Maize plants were subjected to six experimental treatments based on seed priming: three under well-watered conditions (no silicon seed priming and seed priming with 10 and 20 mM sodium silicate solutions) and three corresponding treatments combined with irrigation withdrawal for 15 days to induce drought stress. Morphological traits, biomass accumulation, photosynthetic pigment content, plant water status, and PSI- and PSII-related photochemical parameters were evaluated. Drought stress markedly reduced plant growth, biomass production, relative water content, chlorophyll pigment levels, and photosystem photochemical performance, reflecting a strong negative impact of water deficit on most measured parameters. In particular, root and shoot fresh weights decreased by 75% and 71%, respectively, compared with those of well-watered unprimed control plants, indicating a substantial reduction in biomass accumulation under drought conditions. Furthermore, drought conditions impaired photochemical performance and increased non-regulated energy dissipation, indicative of impaired photosynthetic performance. Silicon seed priming mitigated several drought-induced effects in a trait-dependent manner. Under water-deficit conditions, 20 mM Si produced the strongest improvement in root and shoot fresh weights, whereas 10 mM Si showed stronger responses for selected shoot-growth and PSI-related parameters. Both Si treatments improved leaf water status and photosynthetic stability to varying extents. Collectively, these results indicate that sodium silicate seed priming partially improves drought-related responses in maize seedlings under the conditions of this study by sustaining growth performance, preserving plant water status, and maintaining photosynthetic stability under water-deficit conditions.

Phytoremediation of areas polluted by heavy metals and toxic metalloids is challenging, particularly in arid regions where limited water availability compromises plant establishment. This study shows that several Atriplex species (A. halimus, A. canescens, A. nummularia, A. atacamensis, A. hortensis, etc.) can be used in a phytostabilization and/or phytoextraction strategy. Many of these species are xero-halophytic plants with C4 metabolism that are adapted to harsh conditions. A deep root system allows contaminants to be removed from deep within the soil. These species can accumulate heavy metals and toxic metalloid elements and develop tolerance mechanisms associated with the synthesis of osmoprotective compounds (proline and glycinebetaine), phytochelatins and metallothioneins, and endogenous antioxidant compounds. Sequestration of toxic elements in cell walls or excretion into leaf trichomes contributes to this tolerance. In many cases, transfer factors greater than 1 suggest that these plants can be used to decontaminate polluted sites. Adult plants can produce 3 t.ha-1 of dry matter, and the quantities of pollutants effectively removed from the soil can be significant (from 500 g to several kg per hectare in a single harvest, depending on the pollutant). The application of chelating agents can be useful in increasing the bioavailability of toxic elements, and fertilization, mainly with nitrogen, may be required when the soil is not very fertile and a high plant density (up to 4,000 plants per hectare) is used to help combat erosion. Species of the genus Atriplex are characterized by significant intraspecific genetic variability, and their use therefore requires prior identification of the material best suited to the various pollutants present. Many species of this fascinating genus constitute particularly promising plant material for the low-cost management of large areas of polluted land in arid regions, helping to combat erosion, gradually decontaminate the soil, and restore the ecological balance of marginal areas.

Halophytes, particularly Salicornia species, possess unique metabolic adaptations enabling them to thrive in saline environments, making them valuable for studying plant resilience and phytoremediation. Three extraction techniques (ultrasonication, heat-assisted extraction, and a combination of both) were evaluated, with GC-MS identifying 50 metabolites across all samples, including fatty acids, alkanes, phenols, and spiro compounds. Multivariate analyses (PCA/OPLS-DA) revealed distinct species-specific metabolic profiles with robust statistical performance (R²X = 0.999, R²Y = 1, Q² = 1, CV-ANOVA p = 3.775 × 10⁻¹⁸). These findings underscore divergent adaptive strategies among the three species: S. fruticosa (E1) prioritizes osmotic adjustment and nitrogen storage through amino acid derivatives and pyrroles; in contrast, S. arabica (E2) focuses on robust antioxidant defense and ion balancing, characterized by elevated lactones, sesquiterpenoids, and dicarboxylic acids; and S. perennis (E3) invests in structural integrity, utilizing a high lipophilic content and cuticular waxes as physical barriers against salt and dehydration. Our results highlight these species as effective nature-based biosensors, with S. perennis showing higher capacity for phenol absorption, while S. fruticosa and S. arabica exhibit a stronger affinity for hydrocarbon contaminants, notably spiro[2.4]hepta-4,6-diene, a marker of plastic degradation. This metabolic specialization suggests that each species employs unique biochemical pathways to mitigate saline and xenobiotic stress. Ultimately, this study provides a scientific foundation for using Salicornia species in targeted phytoremediation and environmental monitoring of threatened littoral ecosystems.

Although the physiological mechanisms underlying salt tolerance in wild barley (Hordeum maritimum) are relatively well understood, its recovery after salt stress remains poorly characterized. In this pot experiment, plants were exposed to 200 mM NaCl for 0, 15, 30, 45, or 60 days to evaluate changes in growth and water status. Wild barley exhibited high tolerance during the early stages of salt exposure, maintaining normal growth and water balance for up to 30 days. However, longer exposure reduced plant growth and leaf water content, indicating that prolonged salinity eventually compromises plant performance. To cope with salt stress, plants accumulate soluble sugars, proline, and mineral ions, helping maintain water uptake and cell hydration. Mineral ions, particularly sodium (Na+) and chloride (Cl-), contributed most to osmotic adjustment, accounting for up to 87% of the total osmotic adjustment after 60 days of treatment. Salt stress also induced structural modifications in leaf tissues that enhanced water retention and maintained cell function under saline conditions. Importantly, unlike cultivated barley, H. maritimum fully recovered after the salt treatment ended, restoring both growth and water relations. This strong recovery capacity demonstrates remarkable physiological flexibility and adaptation to changing environmental conditions. These findings highlight the adaptive advantage of wild barley in fluctuating saline environments, where efficient osmotic adjustment, flexible cell wall dynamics, and rapid recovery are key adaptive traits. From both physiological and ecological perspectives, H. maritimum represents a valuable model for studying salt tolerance mechanisms and a promising genetic resource for improving crop salinity resilience.

Soil salinization is a major abiotic stress that impairs plant growth and damages the photosynthetic apparatus. Plant growth promoting rhizobacteria (PGPR) are known to enhance plant tolerance to salinity, despite their mechanistic effects on photosynthesis remain unclear. Therefore, this study investigates the effects of Bacillus subtilis inoculation on the photosynthetic activity of Sulla carnosa under salt stress, with a focus on the functional performance of photosystems I and II. Seeds were either inoculated or not and grown in sterilized soil for one month, before exposure to 0 or 200 mM NaCl salinity for an additional month. Salinity severely reduced shoot and root dry weights by 32

The current investigation explored the potential of Limonium spathulatum extracts as antioxidants for nutritionally enhanced jelly candies. So, the impact of different drying and extraction methods on L. spathulatum phytochemicals was examined, the bioaccessibility of polyphenols was assessed and the impact of extracts incorporation into jelly candies was evaluated. The results revealed that air drying combined with infusion achieved the highest extraction efficiency (114.61

The use of phosphate-solubilizing bacteria (PSB) is a promising strategy to offset the harmful effect of combined salinity and low phosphorus availability and constitutes an affordable solution to enhance agricultural productivity under co-occurring abiotic stresses. In the present study, we investigate the effect of seed inoculation with different PSB isolates on the responses of barley (Hordeum vulgare) seedlings exposed to salt stress, whether individually applied or in combination with phosphorus deficiency. PSB strains used showed beneficial effect by significantly improving barley response under single and/or combined stresses. Yet, effects were strain- and organ-specific. Considering the plant growth promoting effect, GS4f isolate (Pseudomonas sp.) was the most effective strain in relationship with better water status and photosynthesis activity. Seed inoculation with PSB also reduced Na+ content and enhanced K+ content along with higher phosphorus mobilization (as P accumulation and acid phosphatase activity). This was concomitant with decreased H2O2 production resulting in lower MDA content in stressed roots and leaves of inoculated plants. PSB Inoculation triggered the overall plant antioxidant defense, including enzymatic (SOD, CAT and GPX) under simultaneous salinity and low phosphorus availability. Overall, our findings provide valuable information for prospective production of effective biostimulants based on halotolerant PSB and further highlight the possibility of using this promising eco-friendly approach to improve plant growth in P-deficient and salt-affected soils.

The Na⁺/H⁺ antiporter (NHX) gene family in plants encodes proteins that maintain ion homeostasis, particularly under salt stress, by exchanging Na⁺ or K⁺ for H⁺ across cellular membranes. In cultivated barley (Hordeum vulgare L.), NHX genes have been only partially characterized. In this study, we conducted a genome-wide identification of NHX genes in barley, examining their evolutionary relationships, gene structure, and expression patterns under salt stress (200 mM NaCl) in two genotypes differing in salinity tolerance. Seven genes were identified in the H. vulgare genome, and they are unevenly distributed across chromosomes 2 H, 3 H, 4 H, 5 H, and 7 H. Phylogenetic analysis showed that these genes group into three major classes: Vac (HvNHX1, HvNHX2 and HvNHX3), Endo (HvNHX4, HvNHX5 and HvNHX6), and PM (HvNHX7/HvSOS1). Furthermore, exon–intron organization and conserved motif composition were highly conserved within each class. Vac-class NHX proteins were found to contain an amiloride-binding site in TM3 within their N-terminal region. Promoter analysis revealed that HvNHX1, HvNHX5, HvNHX6, and HvNHX7 possess the highest number of abscisic acid (ABA)-responsive elements (ABREs), suggesting potential regulation via the ABA signaling pathway. The protein-protein interaction (PPI) network indicated that several HvNHX proteins interact with HKT, GORK, CHX and KEA partners. Finally, the RT-qPCR analysis revealed a differential expression of NHX genes between the two contrasting barley genotypes in both roots and leaves under salinity. Our findings provide valuable insights into candidate genes that may be targeted in future genetic engineering strategies to enhance salinity tolerance in barley.

Integrating Compost into soil management promotes sustainable nutrient management, strengthens soil resilience, and underpins environmentally friendly agricultural production. This study assessed the interactive effects of compost salinity, application rate (0–40-80-120 t·ha⁻¹), on plant growth and heavy metal allocation in Medicago sativa Gabès variety across different soil textures. The emphasis of the study was on the issue of soil salinization caused by compost application and its effect on plant performance. The accumulation of soluble salts (Na+) and heavy metals was evaluated to in order to understand the interactions between compost, soil and plants affecting nutrient dynamics, sodium toxicity and productivity in saline environments. Although MSW compost enhances soil fertility and crop productivity, its use in arid regions remains uncertain due to elevated soluble salt concentrations that may intensify soil electrical conductivity and hinder plant growth. The substrate’s suitability in degraded land is questionable due to the excessive soluble salts (Sodium Chloride (NaCl), Calcium Sulfate CaSO4, MgSo4…) in MSW that is likely to raise soil salinity and damage plants further. The study found that cumulative biomass increased by 33

Leafy green vegetables highly accumulate cadmium (Cd) in edible tissues, representing a significant health risk for consumers. Despite growing interest in Cd phytotoxicity, studies specifically addressing the interplay between Fe nutrition and Cd stress in lettuce, particularly regarding photosystem integrity and subcellular Cd compartmentalization, remain limited. This study evaluates the effectiveness of iron supplementation to reduce Cd accumulation in lettuce (Lactuca sativa L.), based on the hypothesis of antagonistic interactions between Cd and Fe. Lettuce plants were hydroponically cultivated using 10, 50, or 250 µM Fe, with or without cadmium (15 µM CdCl2). Plant growth, photosynthetic performance, and cadmium accumulation were assessed. An integrated biophysical approach was applied to evaluate photosystem I and II functional integrity, and subcellular Cd distribution among soluble, cell wall, and organelle fractions was determined to provide mechanistic insights into Fe-mediated Cd tolerance. Cd exposure induced leaf chlorosis and strongly inhibited plant growth, reducing shoot and whole-plant dry weight by 42

Despite their common co-existence in soil, the combined toxic effects of salinity and cadmium on plants is largely unexplored. In this study, a hydroponic experiment was conducted to determine the effect of long-term exposure of two barley genotypes differing in their salinity tolerance; Rihane (RH, salt tolerant) and Lemsi (LM, salt sensitive) to individual NaCl and Cd and their combination. Results showed that salinity alone or in combination with Cd significantly reduced growth, photosynthetic rates (A, E, and gs), leaf pigments and nutrient elements (K, Fe, Mg, and Zn) amounts. Meanwhile, it increased Na and Ca amounts, electrical conductivity, proline, and carotenoid levels. The overall effect was greater under the sole NaCl application in LM genotype than in RH. In contrast, the sole Cd application slightly reduced growth parameters. LM demonstrated higher amounts of Cd in roots (44.038 mu g Plant-1) and shoots (40.8 mu g Plant-1) compared to RH (39.8 and 21.5 mu g Plant-1, respectively). Salinity-Cd co-exposure reduced total Cd content by 65% in LM and 61% in RH, which could be due to an enhanced competition between Na and Cd. This was accompanied by elevated levels of proline and carotenoids as well as the upregulation of genes mediating heavy metal transport (HvZIP8, HvABCG25, and HvAHA1). A substantial role of nonselective cation channels (NSCCs) involved in Cd transport that may be inhibited by NaCl and other essential elements like Ca was noted. Overall, HvZIP8, HvAHA1 and HvABCG25 transporters could be used in future biotechnological applications to reduce Cd accumulation in barley.

Sea barley ( Hordeum marinum ), a halophyte among the largely glycophytic Hordeum species, is a valuable genetic resource for elucidating salt tolerance mechanisms in cereals. This study assessed the physiological, biochemical, and antioxidant responses of two H. marinum genotypes (AR and SH) and the salt‐tolerant Hordeum vulgare L. cv. Rihane (RH) under 400 mM NaCl for 3 weeks. Results revealed a marked growth reduction in RH, correlating with higher accumulation of Na and Cl in leaves (six‐ and four‐fold, respectively) and roots (2.29‐ and 1.67‐fold, respectively) compared to AR and SH. In contrast, SH and AR maintained higher leaf osmotic potential (−1.96 and −2.11 MPa, respectively) and better root K acquisition under salinity. Despite an increase in leaf proline content and antioxidant enzyme activities, RH displayed elevated hydrogen peroxide (H 2 O 2 ) and malondialdehyde (MDA) accumulation compared to SH and AR, indicating insufficient antioxidant defenses against salt‐induced oxidative stress. Correlation analysis revealed positive associations between Cl, Na, H 2 O 2 , and MDA, while negative correlations emerged with K, osmotic potential, and antioxidant enzyme activities. The superior salt tolerance of AR and SH was closely linked to their ability to maintain osmotic balance, regulate ion homeostasis, and mitigate oxidative stress. These traits offer promising targets for improving salt tolerance in cultivated barley and other cereals.

Soil contamination by heavy metals, such as zinc, has significant environmental consequences. Phytoremediation, among various remediation techniques, has been developed and applied to restore contaminated soils. However, phytoremediation has limitations, especially related to slow plant growth. This issue is often influenced by the toxicity of contaminants. This study aims to investigate the interaction between zinc and silicon in oleander (Nerium oleander L.). Specifically, it examines the impact on growth, photosynthetic and biochemical behaviors, as well as the plant's phytoremediation capacity. Oleander plants were exposed to four treatments. These included two adequate zinc treatments (0.76 μM Zn) combined with two silicon concentrations (0 mM and 0.5 mM Si). Additionally, two zinc-toxic treatments (1800 μM Zn) were applied with the same silicon concentrations. The results demonstrated that zinc toxicity had detrimental effects on most of the measured parameters. However, the negative impacts were substantially alleviated by silicon supplementation. The application of silicon reduced leaf chlorosis and enhanced biomass production. It also increased photosynthetic pigment content, improved gas exchange, and maintained the oxidation state of photosystem I (PSI). Additionally, it preserved membrane integrity. Moreover, silicon influenced soluble protein levels and increased antioxidant enzyme activities. This was especially true for guaiacol peroxidase, which helped mitigate oxidative stress. Overall, silicon had beneficial effects on the phytoremediation capacity of N. oleander. Therefore, fertilization rich in silicon could represent an effective solution for enhancing the phytoremediation capacity of this species, minimizing one of the major disadvantages of phytoremediation, namely low biomass production influenced by toxicity.

Atriplex halimus L. is a promising xero-halophyte species for phytoremediation purposes but displays high levels of genetic variability. As an attempt to select uniform material suitable for phytomanagement, five clones were established by cuttings from three non-polluted sites (Tunis, Nabeul and Sfax), one moderately Pb-polluted site (Sousse) and a highly polymetallic polluted mining site (Gafsa). Cuttings were cultivated during 90 days under controlled conditions on soil issued from the most polluted area. The clone from Gafsa accumulated higher concentrations of metals in roots (Cr) and leaves (Cd, Sr, Zn and Cu) than other clones. Gafsa showed the highest absorption efficiency, translocation factor, bioconcentration factor and bioaccumulation coefficient compared to other clones but displayed the lowest relative growth rate (RGR) value while the highest RGR was found in the clone from Sousse. Heavy metal tolerance in Gafsa was not related to a more efficient management of oxidative stress or higher concentration of phytochelatins. Total amount of Sr and Zn removed from the substrate was the highest for Sousse while removal of Cd, Cu, Cr and Ni was the highest for Gafsa. It is concluded that cuttings allow to obtain uniform material for phytoremediation by Atriplex halimus and that combining different clones with complementary properties is an attractive option for phytomanagement of polymetallic polluted soils by this species.

Hydroponic cultivation offers a promising solution to water scarcity by using less water than traditional soil-based agriculture. Although the integration of silicon (Si) in hydroponic systems is still limited, its foliar application is crucial for optimizing plant growth, enhancing resilience, and improving productivity. Therefore, this study aims to explore the effects of silicon foliar application on growth, yield parameters, and photosynthetic activity of one of the most important fruit vegetables worldwide—tomatoes (Solanum lycopersicum L.)—cultivated in a closed hydroponic system. Tomato plants were subjected to a weekly silicon foliar spray (1 mM Na2SiO3) over a period of 10 weeks. Our results demonstrate that silicon foliar spray in hydroponically grown tomatoes significantly improved photosynthetic pigment concentration and enhanced the photochemical efficiency of the photosystems, particularly the quantum yields Y(I) and Y(II). Moreover, silicon treatment resulted in reduced energy dissipation within the photosystems, as indicated by decreases in Y(NPQ), Y(NO), and Y(ND), along with enhanced oxidation of PSI (P700ox). These physiological improvements were directly linked to increased stem width and length, and a substantial boost in plant dry weight compared with untreated controls. Consequently, the silicon foliar spray resulted in a significant increase in the number of racemes, flowers, and fruits. Ultimately, these enhancements culminated in a 69% increase in fruit biomass yield (r2 = 0.70; p < 0.01), highlighting the positive impact of silicon on tomato productivity in hydroponic systems. These findings suggest that silicon foliar application optimizes growth and photosynthetic efficiency while reducing energy dissipation. Consequently, silicon supplementation enhances tomato productivity in hydroponic systems, offering a promising solution for increasing yields in water-efficient agriculture.

Cadmium (Cd) contamination represents a significant environmental challenge, posing serious risks to human health. This study aims to examine the relationship between two trace metals (zinc (Zn) and cadmium (Cd)) in Lactuca sativa, a leafy vegetable known for its propensity to accumulate heavy metals. Two weeks after germination, lettuce seedlings were subjected to Cd treatments (0 and 10 µM), in combination with varying Zn concentrations (1, 10, and 100 µM). The efficacy of Zn supplementation in alleviating Cd toxicity was notably observed in the shoots, where partial to full recovery of key photosynthetic parameters, including transpiration rate, water use efficiency, maximum quantum efficiency (Fv/Fm), and maximum operating efficiency of PSII (Fv'/Fm'), was documented. Additionally, the efficiency of Zn supply in restricting Cd increase in aerial parts was obvious, particularly at the highest Zn concentration (100 µM), which coincided with increased Zn accumulation. By contrast, at this level, root Cd absorption was enhanced, accompanied by a marked inhibition of Zn uptake. These results indicate an antagonistic effect linking these two heavy metals, likely due to competitive interactions for shared membrane transporters. Overall, the findings suggest that appropriate Zn supplementation may serve as a viable agronomic strategy to mitigate Cd accumulation in species grown in lands contaminated with Cd, thereby offering a potential solution for improving food safety and environmental health.

The goal of the current study was to investigate the effects of seed priming with salicylic acid (SA) on seed germination parameters, seedling growth traits, nutritional element mobilization, and oxidative stress status in two barley species that were subjected to various salt treatments. The findings demonstrated that salinity reduced a number of germination parameters in unprimed seeds and impacted seedling growth by impeding both species’ necessary nutrient mobilization. Under this abiotic stress, a noticeable rise in malondialdehyde and electrolyte leakage was also noted. Interestingly, pretreating seeds with SA improved seed germination and seedling growth performance under either 100 mM or 200 mM NaCl treatments. In fact, SA improved the length and dry weight of stressed seedlings of both barley species in addition to increasing the germination rate and mean daily germination. Additionally, SA increased the content of calcium, iron, magnesium, and potassium while lowering the concentrations of sodium and malondialdehyde and electrolyte leakage. It is significant to note that, in comparison to Hordeum maritimum, the positive effects of this hormone were more noticeable in stressed Hordeum vulgare species.

Soil pollution emerging from mining activities is a worldwide environmental concern. Finding biological solutions to this problem is of paramount importance in reducing metal toxicity and limiting its spread. In this context, halophytes-assisted phytoremediation has been widely considered as a promising alternative to remediate both salt-affected and heavy metal-contaminated soils. In this study, three halophytes Sesuvium portulacastrum, Carpobrotus edulis and Mesembryanthemum cordifolium have been evaluated for their metal phytoextraction potential performed on a mining soil collected from the mining phosphate basin of Gafsa. A 7-month greenhouse pot experiment was conducted under two cultivation systems: mono-cropping and co-cropping. The effect of co-cropping on the phytoextraction efficiency was highlighted and the associated phytoremediation indexes were assessed. Results showed that the crude collected soil revealed strong Cd and Sr contamination, moderate Cr and Ni levels, and low Cu and Zn concentrations. Compared to mono-cropping, co-cropping enhanced Cd uptake efficiency achieving a 40

Salt accumulation can degrade soil properties, decrease its productivity, and harm its ecological functions. Introducing salt-tolerant plant species associated with arbuscular mycorrhizal fungi (AMF) can act as an effective biological method for restoring salinized soils. AMF colonize plant roots and improve their nutrient acquisition capacity. However, there is limited knowledge on how AMF affects the production of signaling molecules, e.g., abscisic acid (ABA), salicylic acid (SA), and jasmonic acid (JA), related to plant–microbe interactions under salinity. Here, we assess the potential benefits of the AMF Rhizophagus intraradices in enhancing plant growth and nutrient uptake in addition to modulating stress hormone signaling levels (ABA, SA, and JA) of the facultative halophyte Sulla carnosa under saline conditions. Plants were grown in pots filled with soil and irrigated with 200 mM NaCl for 1 month. AMF symbiosis substantially increased the shoot dry weight (+107%), root dry weight (+67%), photosynthetic pigment content (chlorophyll a, chlorophyll b, and carotenoids), and nutrient uptake (C, N, P, K, Cu, and Zn) while significantly limiting the increase in the shoot Na+ concentration and H2O2 content caused by salinity stress. Mycorrhizal symbiosis significantly enhanced the root and shoot SA levels by 450% and 32%, respectively, compared to the stressed non-inoculated plants, potentially contributing to enhanced systemic resistance and osmotic adjustment under saline conditions. Salt stress increased the shoot ABA content, especially in R. intraradices-inoculated plants (113% higher than in stressed non-mycorrhizal plants). These findings confirm that AMF mitigated the adverse effects of salinity on S. carnosa by increasing the SA and ABA levels and reducing oxidative damage.