Nanomaterial sulfonated graphene oxide advances the tolerance against nitrate and ammonium toxicity by regulating chloroplastic redox balance, photochemistry of photosystems and antioxidant capacity in Triticum aestivum.

The current study was designed to assess nanomaterial sulfonated graphene oxide (SGO) potential in improving tolerance of wheat chloroplasts against nitrate (NS) and ammonium (AS) toxicity. Triticum aestivum cv. Ekiz was grown under SGOs (50-250-500 mg L-1) with/without 140 mM NS and 5 mM AS stress. SGOs were eliminated the adverse effects produced by stress on chlorophyll fluorescence, potential photochemical efficiency and physiological state of the photosynthetic apparatus. SGO reversed the negative effects on these parameters. Upon SGOs exposure, the induced expression levels of photosystems-related reaction center proteins were observed. SGOs reverted radical accumulation triggered by NS by enabling the increased superoxide dismutase (SOD) activity and ascorbate (AsA) regeneration. Under AS, the turnover of both AsA and glutathione (GSH) was maintained by 50-250 mg L-1 SGO by increasing the enzymes and non-enzymes related to AsA-GSH cycle. 500 mg L-1 SGO prevented the radical over-accumulation produced by AS via the regeneration of AsA and peroxidase (POX) activity rather than GSH regeneration. 50-250 mg L-1 SGO protected from the NS+AS-induced disruptions through the defense pathways connected with AsA-GSH cycle represented the high rates of AsA/DHA and, GSH/GSSG and GSH redox state. Our findings specified that SGO to NS and AS-stressed wheat provides a new potential tool to advance the tolerance mechanism.

Naringenin induces tolerance to salt/osmotic stress through the regulation of nitrogen metabolism, cellular redox and ROS scavenging capacity in bean plants.

The present study was conducted to uncover underlying possible effect mechanisms of flavonoid naringenin (Nar, 0.1-0.4 mM) in nitrogen assimilation, antioxidant response, redox status and the expression of NLP7 and DREB2A, on salt (100 mM NaCl) and osmotic-stressed (10% Polyethylene glycol, -0.54 MPa) Phaseolus vulgaris cv. Yunus 90). Nar ameliorated salt/osmotic stresses-induced growth inhibition and improved the accumulation of proline, glycine betaine and choline. In response to stress, Nar increased endogenous content of nitrate (NO3-) and nitrite (NO2-) by regulating of nitrate reductase and nitrite reductase. Stress-triggered NH4+ was eliminated with Nar through increases in glutamine synthetase and glutamate synthase. After NaCl or NaCl + PEG exposure, Nar utilized the aminating activity of glutamate dehydrogenase in the conversion of NH4+. The stress-inducible expression levels of DREB2A were increased further by Nar, which might have affected stress tolerance of bean. Nar induced effectively the relative expression of NLP7 in the presence of the combination or alone of stress. Also, the impaired redox state by stress was modulated by Nar and hydrogen peroxide (H2O2) and TBARS decreased. Nar regulated the different pathways for scavenging of H2O2 under NaCl and/or PEG treatments. When Nar + NaCl exposure, the damage was removed by superoxide dismutase (SOD), catalase (CAT), POX (only at 0.1 mM Nar + NaCl) and AsA-GSH cycle. Under osmotic stress plus Nar, the protection was manifested by activated CAT and, glutathione S-transferase and the regeneration of ascorbate. 0.1 mM Nar could protect bean plant against salt/osmotic stresses, likely by regulating nitrogen assimilation pathways, improving expression levels of genes associated with tolerance mechanisms and modulating the antioxidant capacity and AsA-GSH redox-based systems.

Flavonoid Naringenin Alleviates Short-Term Osmotic and Salinity Stresses Through Regulating Photosynthetic Machinery and Chloroplastic Antioxidant Metabolism in Phaseolus vulgaris.

The current study was conducted to demonstrate the possible roles of exogenously applied flavonoid naringenin (Nar) on the efficiency of PSII photochemistry and the responses of chloroplastic antioxidant of salt and osmotic-stressed Phaseolus vulgaris (cv. Yunus90). For this aim, plants were grown in a hydroponic culture and were treated with Nar (0.1 mM and 0.4 mM) alone or in a combination with salt (100 mM NaCl) and/or osmotic (10% Polyethylene glycol, -0.54 MPa). Both caused a reduction in water content (RWC), osmotic potential (ΨΠ), chlorophyll fluorescence (Fv/Fm), and potential photochemical efficiency (Fv/Fo). Nar reversed the changes on these parameters. The phenomenological fluxes (TRo/CS and ETo/CS) altered by stress were induced by Nar and Nar led to a notable increase in the performance index (PIABS) and the capacity of light reaction [ΦPo/(1-ΦPo)]. Besides, Nar-applied plants exhibited higher specific fluxes values [ABS/RC, ETo/RC, and ΨEo/(1-ΨEo)] and decreasing controlled dissipation of energy (DIo/CSo and DIo/RC). The transcripts levels of psbA and psbD were lowered in stress-treated bean but upregulated in Nar-treated plants after stress exposure. Nar also alleviated the changes on gas exchange parameters [carbon assimilation rate (A), stomatal conductance (gs), intercellular CO2 concentrations (Ci), transpiration rate (E), and stomatal limitation (Ls)]. By regulating the antioxidant metabolism of the isolated chloroplasts, Nar was able to control the toxic levels of hydrogen peroxide (H2O2) and TBARS (lipid peroxidation) produced by stresses. Chloroplastic superoxide dismutase (SOD) activity reduced by stresses was increased by Nar. In response to NaCl, Nar increased the activities of ascorbate peroxidase (APX), glutathione reductase (GR), monodehydroascorbate reductase (MDHAR), and dehydroascorbate reductase (DHAR), as well as peroxidase (POX). Nar protected the bean chloroplasts by minimizing disturbances caused by NaCl exposure via the ascorbate (AsA) and glutathione (GSH) redox-based systems. Under Nar plus PEG, Nar maintained the AsA regeneration by the induction of MDHAR and DHAR, but not GSH recycling by virtue of no induction in GR activity and the reduction in GSH/GSSG and GSH redox state. Based on these advances, Nar protected in bean chloroplasts by minimizing disturbances caused by NaCl or PEG exposure via the AsA or GSH redox-based systems and POX activity.

Rare-earth element scandium improves stomatal regulation and enhances salt and drought stress tolerance by up-regulating antioxidant responses of Oryza sativa.

Oryza sativa L. cv. Gönen grown in hydroponic culture was treated with scandium (Sc; 25 and 50 µM) alone or in combination with salt (100 mM NaCl) and/or drought (5% PEG-6000). Stress caused a decrease in growth (RGR), water content (RWC), osmotic potential (ΨΠ), chlorophyll fluorescence (Fv/Fm) and potential photochemical efficiency (Fv/Fo). Sc application prevented the decreases of these parameters. Sc also alleviated the changes on gas exchange parameters (carbon assimilation rate (A), stomatal conductance (gs), intercellular CO2 concentrations (Ci), transpiration rate (E) and stomatal limitation (Ls)). Stress caused no increase in superoxide dismutase (SOD) activity. After induvial applied NaCl or PEG, catalase (CAT) and ascorbate peroxidase (APX) showed an enhancement in activation and tried to scavenge of hydrogen peroxide (H2O2). On the other hand, in plants with the combination form of NaCl and PEG, only CAT activity was induced. Sc applications to NaCl-treated rice led to an increase of SOD, APX, glutathione reductase (GR), monodehydroascorbate reductase (MDHAR) and dehydroascorbate reductase (DHAR) as well as peroxidase (POX). Sc under NaCl could be maintained both ascorbate (AsA) and glutathione (GSH) regeneration. Despite of induction of MDHAR and DHAR under Sc plus PEG, Sc did not maintain AsA redox state because of no induction in APX activity. However, GSH pool could be regenerated by induction in DHAR and GR in this group. Sc application (especially for 25 µM) in rice exposed to NaCl + PEG resulted an enhancement in APX and MDHAR and so Sc could be partially provided AsA regeneration. Since no increases in DHAR and GR were observed, GSH pool was reduced. Due to this activation of antioxidant enzymes, stress-induced H2O2 and TBARS content (lipid peroxidation) significantly decreased in rice with Sc applications. Sc in plants with stress also increased the transcript levels of OsCDPK7 and OsBG1 related to stomatal movement and signaling pathway. Consequently, Sc protected the rice plants by minimizing disturbances caused by NaCl or PEG exposure via the AsA-GSH redox-based systems.

Hydrogen sulfide (H2S) and nitric oxide (NO) alleviate cobalt toxicity in wheat (Triticum aestivum L.) by modulating photosynthesis, chloroplastic redox and antioxidant capacity.

The role of hydrogen sulfide (H2S)/nitric oxide (NO) in mitigating stress-induced damages has gained interest in the past few years. However, the protective mechanism H2S and/or NO has towards the chloroplast system through the regulation of redox status and activation of antioxidant capacity in cobalt-treated wheat remain largely unanswered. Triticum aestivum L. cv. Ekiz was treated with alone/in combination of a H2S donor (sodium hydrosulfide (NaHS,600µM)), a NO donor (sodium nitroprusside (SNP,100µM)) and a NO scavenger (rutin hydrate (RTN,50µM)) to assess how the donors affect growth, water relations, redox and antioxidant capacity in chloroplasts, under cobalt (Co) concentrations of 150-300 µM. Stress decreased a number of parameters (growth, water content (RWC), osmotic potential (ΨΠ), carbon assimilation rate, stomatal conductance, intercellular CO2 concentrations, transpiration rate and the transcript levels of rubisco, which subsequently disrupt the photosynthetic capacity). However, SNP/NaHS counteracted the negative effects of stress on these aforementioned parameters and RTN application with stress/non-stress was reversed these effects. Hydrogen peroxide (H2O2) and TBARS were induced under stress in spite of activated ascorbate peroxidase (APX). SNP/NaHS under stress increased activation of superoxide dismutase (SOD), peroxidase (POX), APX, glutathione reductase (GR), monodehydroascorbate reductase (MDHAR), dehydroascorbate reductase (DHAR), ascorbate (tAsA) and glutathione (GSH). In conclusion, NaHS/SNP are involved in the regulation and modification of growth, water content, rubisco activity and up-regulation of ascorbate-glutathione cycle (AsA-GSH) in chloroplast under stress.

The humic acid-induced changes in the water status, chlorophyll fluorescence and antioxidant defense systems of wheat leaves with cadmium stress.

The using of bio-stimulant in plants grown under stress conditions for enhancing nutrition efficiency and crop quality traits is an effective approach. One of the bio-stimulants, humus material, is defined as humic acid (HA). HA application as a promotion of plant growth to plants grown in the heavy metals-contaminated soils has promised hope in terms of effects on plants but the its limiting effect is the application dose. Therefore, the wheat seedlings were grown in hydroponic culture for 21 d and the various concentrations of humic acid (HA; 750 or 1500 mg L-1) were treated alone or in combination with cadmium (Cd) stress (100 or 200 µM) for 7 d. The results showed that after Cd stress treatment, water content (RWC), osmotic potential (ΨΠ) and chlorophyll fluorescence parameters decreased and proline content (Pro) increased for 7 d. In spite of activated peroxidase (POX) and ascorbate peroxidase (APX), stress induced the toxic levels of hydrogen peroxide (H2O2) accumulation. Cd stress triggered lipid peroxidation (TBARS content). HA application successfully eliminated the negative effects of stress on RWC, ΨΠ and photosynthetic parameters. In the presence of HA under stress, the increased activation of superoxide dismutase (SOD), catalase (CAT) and NADPH-oxidase (NOX) enzymes and ascorbate, glutathione and GSH/GSSG ratio observed. Only 750 mg L-1 HA under stress conditions induced the activities of monodehydroascorbate reductase (MDHAR) and dehydroascorbate reductase (DHAR), and dehydroascorbate (DHA) content. After the combined application of HA and Cd stress, the low contents of H2O2 and TBARS maintained in wheat leaves. Hence, HA successfully eliminated the toxicity of Cd stress by modulating the water status, photosynthetic apparatus and antioxidant activity in wheat leaves.

Upregulation of antioxidant enzymes by exogenous gallic acid contributes to the amelioration in Oryza sativa roots exposed to salt and osmotic stress.

The aim of this study is to elucidate the influence of the exogenous application of gallic acid (GLA) in alleviating the detrimental effects of salinity (NaCl), osmotic stress (polyethylene glycol; PEG), and their combination in Oryza sativa L. roots. To produce same osmotic potential (-0.5 MPa), 3-week-old rice seedlings were treated with 120 mM NaCl and/or 20 % PEG6000 with/without GLA (0.75 and 1.5 mM) treatments for 72 h. Both alone and combination of stresses decreased growth (RGR) and osmotic potential (Ψ Π). Moreover, stress caused a significant increase in proline (Pro) and hydrogen peroxide (H2O2) contents. Also, Pokkali and IR-28 had higher H2O2-scavenging enzyme activities including catalase (CAT), ascorbate peroxidase (APX), and glutathione reductase (GR) activities in NaCl-treated roots. Only CAT activity was induced in both cultivars with PEG. Therefore, the enhanced levels of lipid peroxidation (thiobarbituric acid reactive substances (TBARS)) were more pronounced under PEG than NaCl. However, GLA significantly mitigated NaCl and/or PEG-induced stress injury. Under salinity, TBARS was lesser in GLA-applied rice that was associated with greater activities of superoxide dismutase (SOD), peroxidase (POX), and APX. GLA in the presence of PEG improved the activities of CAT and POX. According to these findings, GLA alleviated the damaging effects of NaCl and/or PEG (especially under NaCl) by improving the antioxidative system in rice. This is the first study elucidating the effects of GLA on tolerance to salinity, osmotic stress, and their combination in plants.

The role of antioxidant responses on the tolerance range of extreme halophyte Salsola crassa grown under toxic salt concentrations.

Salsola crassa (Amaranthaceae) is an annual halophytic species and naturally grows in arid soils that are toxic to most plants. In order to study the effects of salinity on their antioxidant system and to determine the tolerance range against salt stress, S. crassa seeds were grown with different concentrations of NaCl (0, 250, 500, 750, 1000, 1250 and 1500mM) for short (15d) and long-term (30d). Results showed that growth (RGR), water content (RWC) and osmotic potential (ΨΠ) decreased and, proline content (Pro) increased at prolonged salt treatment. Unlike K(+) and Ca(2+) contents, S. crassa highly accumulated Na(+) and Cl(-) contents. Chlorophyll fluorescence (Fv/Fm) only decreased in response to 1500mM NaCl at 30d. No salt stimulation of superoxide anion radical (O2(•-)) content was observed in plants treated with the range of 0-500mM NaCl during the experimental period. NaCl increased superoxide dismutase (SOD) activity depending on intensities of Mn-SOD and Fe-SOD isozymes except in 1500mM NaCl-treated plants at 30d. In contrast to catalase (CAT), peroxidase (POX) activity increased throughout the experiment. Also, salinity caused an increase in glutathione reductase (GR) and glutathione peroxidase (GPX) and decreased in ascorbate peroxidase (APX), monodehydroascorbate reductase (MDHAR) and dehydroascorbate reductase (DHAR) at 15d. Both total ascorbate (tAsA) and glutathione (tGlut) contents significantly increased in treated plants with 1000-1500mM NaCl at 15d. After 0-1000mM NaCl stress, H2O2 and TBARS contents were similar to control groups at 15d, which were consistent with the increased antioxidant activity (POX, GR and GPX). However, H2O2 content was more pronounced at 30d. Therefore, S. crassa exhibited inductions in lipid peroxidation (TBARS content) in response to extreme salt concentrations. These results suggest that S. crassa is tolerant to salt-induced damage at short-term treatments as well as extreme salt concentrations.

Variations in osmotic adjustment and water relations of Sphaerophysa kotschyana: Glycine betaine, proline and choline accumulation in response to salinity.

BACKGROUND: Sphaerophysa kotschyana Boiss. is naturally distributed in overly salty regions. The key to the completion of the life cycles of S. kotschyana in harsh saline soils may be hidden in changes of its osmo-protectants, but there is currently no information about the interaction between osmotic adjustment and water relations in adaptation to saline conditions. The aim of this article was to determine growth, relative growth rate (RGR), relative water content (RWC), osmotic potential (ΨΠ), photosynthetic efficiency (Fv/Fm), thiobarbituric acid-reactive substances (TBARS) and osmo-protectant contents [proline (Pro), choline (Cho) and glycine betaine (GB)] in S. kotschyana leaves and roots exposed to 0, 150 or 300 mM NaCl for 7 and 14 d (days). RESULTS: The results clearly showed that the reductions in growth, RWC, Fv/Fm, RGR and ΨΠ were more pronounced at 300 mM, especially after 14 d. In the same group, the highest increase in TBARS was recorded in roots (126%) and leaves (31%). The induction at 150 mM was not as high. Therefore, roots appear to be the most vulnerable part of this plant. Moreover, S. kotschyana was able to withstand short-term low salinity. CONCLUSIONS: The osmo-protectant accumulation in S. kotschyana as a salinity acclimation or adaptation was sufficient for toleration of low salt concentration (150 mM). In contrast, the plants exposed to the highest NaCl concentration (300 mM) were not able to maintain the ability to prevent water loss because of further decrease in root/shoot ratio of fresh weight (FW) and dry weight (DW), RWC and RGR.

Sphaerophysa kotschyana, an endemic species from Central Anatolia: antioxidant system responses under salt stress.

Sphaerophysa kotschyana is a Turkish endemic and endangered plant that grows near Salt Lake, in Konya, Turkey. However, little is known about the ability of this plant to generate/remove reactive oxygen species (ROS) or its adaptive biochemical responses to saline environments. After exposure of S. kotschyana to 0, 150, and 300 mM NaCl for 7 and 14 days, we investigated (1) the activities and isozyme compositions of the antioxidant enzymes superoxide dismutase (SOD), catalase (CAT), peroxidase (POX), ascorbate peroxidase (APX), and glutathione reductase (GR); (2) the oxidative stress parameters NADPH oxidase (NOX) activity, lipid peroxidation (MDA), total ascorbate (tAsA) content, and total glutathione content (tGlut); and (3) ROS levels for superoxide anion radical (O 2 (·-) ), hydrogen peroxide (H2O2), hydroxyl radicals (OH·), and histochemical staining of O 2 (·-) and H2O2. H2O2 content increased after 14 days of salt stress, which was consistent with the results from histochemical staining and NOX activity measurements. In contrast, oxidative stress induced by 150 mM NaCl was more efficiently prevented, as indicated by low malondialdehyde (MDA) levels and especially at 7 days, by increased levels of SOD, POX, APX, and GR. However, at 300 mM NaCl, decreased levels of protective enzymes such as SOD, CAT, POX, and GR, particularly with long-term stress (14 days), resulted in limited ROS scavenging activity and increased MDA levels. Moreover, at 300 mM NaCl, the high H2O2 content caused oxidative damage rather than inducing protective responses against H2O2. These results suggest that S. kotschyana is potentially tolerant to salt-induced damage only at low salt concentrations.