The emerging key role of reactive sulfur species in abiotic stress tolerance in plants.

In plants, sulfur plays a critical role in the formation of important biomolecules such as cysteine, methionine, and tripeptide glutathione. Thiol groups, composed of sulfur, are essential to numerous metabolic processes. The easy and reversible oxidation and reduction of thiol groups have drawn attention to the redox regulation of cellular metabolism. Reactive sulfur species (RSS), including hydrogen sulfide (H2 S), persulfides, and polysulfides, are synthetized in all living organisms, mainly from cysteine, and have been recognized in the last two decades as very important molecules in redox regulation. RSS are considered potent signaling molecules, being involved in the regulation of virtually all aspects of cell function. With regard to stress, reactive species and the antioxidant machinery maintain a delicate balance that gets disturbed under stress conditions, wherein reactive species biosynthesis, transportation, scavenging, and overall metabolism become decisive for plant survival. While reactive oxygen and nitrogen species have been much discussed over recent years, research into RSS biosynthesis, signaling, and relation to abiotic stresses is still nascent. RSS evolved long before reactive oxygen species, and because both are metabolized by catalase, it has been suggested that "antioxidant" enzymes originally evolved to regulate RSS and may still do so today. In this review, we have tried to summarize the generation, signaling, and interaction of RSS in plant systems and to discuss in detail the roles under various abiotic stresses.

Involvement of ethylene in melatonin-modified photosynthetic-N use efficiency and antioxidant activity to improve photosynthesis of salt grown wheat.

The involvement of melatonin in the regulation of salt stress acclimation has been shown in plants in this present work. We found that the GOAL cultivar of wheat (Triticum aestivum L.) was the most salt-tolerant among the investigated cultivars, GOAL, HD-2967, PBW-17, PBW-343, PBW-550, and WH-1105 when screened for tolerance to 100 mM NaCl. The application of 100 µM melatonin maximally reduced oxidative stress and improved photosynthesis in the cv. GOAL. Melatonin supplementation reduced salt stress-induced oxidative stress by upregulating the activity of antioxidant enzymes, such as superoxide dismutase (SOD), ascorbate peroxidase (APX), and glutathione reductase (GR), and reduced the glutathione (GSH) production. This resulted in increased membrane stability, photosynthetic-N use efficiency and photosynthesis in plants. The application of 50 µM of the ethylene biosynthesis inhibitor aminoethoxyvinylglycine (AVG) in the presence of melatonin and salt stress increased H2 O2 content but reduced GR activity and GSH, photosynthesis, and plant dry mass. This signifies that melatonin-mediated salt stress tolerance was related to ethylene synthesis as it improved antioxidant activity and photosynthesis of plants under salt stress. Thus, the interaction of melatonin and ethylene bears a prominent role in salt stress tolerance in wheat and can be used to develop salt tolerance in other crops.

Exogenously-Sourced Ethylene Positively Modulates Photosynthesis, Carbohydrate Metabolism, and Antioxidant Defense to Enhance Heat Tolerance in Rice.

The effect of exogenously-applied ethylene sourced from ethephon (2-chloroethyl phosphonic acid)was studied on photosynthesis, carbohydrate metabolism, and high-temperature stress tolerance in Taipei-309 and Rasi cultivars of rice (Oryza sativa L.). Heat stress increased the content of H2O2 and thiobarbituric acid reactive substances (TBARS)more in Rasi than Taipei-309. Further, a significant decline in sucrose, starch, and carbohydrate metabolism enzyme activity and photosynthesis was also observed in response to heat stress. The application of ethephon reduced H2O2 and TBARS content by enhancing the enzymatic antioxidant defense system and improved carbohydrate metabolism, photosynthesis, and growth more conspicuously in Taipei-309 under heat stress. The ethephon application enhanced photosynthesis by up-regulating the psbA and psbB genes of photosystem II in heat-stressed plants. Interestingly, foliar application of ethephoneffectively down-regulated high-temperature-stress-induced elevated ethylene biosynthesis gene expression. Overall, ethephon application optimized ethylene levels under high-temperature stress to regulate the antioxidant enzymatic system and carbohydrate metabolism, reducing the adverse effects on photosynthesis. These findings suggest that ethylene regulates photosynthesis via carbohydrate metabolism and the antioxidant system, thereby influencing high-temperature stress tolerance in rice.

Hydrogen peroxide potentiates defense system in presence of sulfur to protect chloroplast damage and photosynthesis of wheat under drought stress.

The involvement of hydrogen peroxide (H2 O2 ) combined with sulfur (S) was studied in the protection of the photosynthetic performance of wheat (Triticum aestivum L.) under drought stress. The mechanisms of S-assimilation, the activity of antioxidants, glucose sensitivity, water and osmotic relations and abscisic acid (ABA) content were the focus. The combined application of 50 µM H2 O2 and 100 mg S kg-1 soil (sulfur) resulted in a marked increase in S-assimilation and activity of antioxidant enzymes, with decreased glucose sensitivity and ABA content causing improvement in the structure and function of the photosynthetic apparatus under drought stress. The photosynthetic performance, pigment system (PS) II activity, and growth were improved conspicuously by H2 O2 in the presence of S, as H2 O2 induced S-assimilation capacity, the activity of antioxidant enzymes, and GSH synthesis under drought stress. Our study shows that H2 O2 is more effective in the reversal of drought stress in the presence of S through its influence on S-assimilation, glucose sensitivity, and antioxidant system. These results provide evidence for the effectiveness of H2 O2 in improving photosynthesis under drought stress in the presence of S.

Crosstalk between abscisic acid and nitric oxide under heat stress: exploring new vantage points.

Heat stress adversely affects plants growth potential. Global warming is reported to increase in the intensity, frequency, and duration of heatwaves, eventually affecting ecology, agriculture and economy. With an expected increase in average temperature by 2-3 °C over the next 30-50 years, crop production is facing a severe threat to sub-optimum growth conditions. Abscisic acid (ABA) and nitric oxide (NO) are growth regulators that are involved in the adaptation to heat stress by affecting each other and changing the adaptation process. The interaction between these molecules has been discussed in various studies in general or under stress conditions; however, regarding high temperature, their interaction has little been worked out. In the present review, the focus is shifted on the role of these molecules under heat stress emphasizing the different possible interactions between ABA and NO as both regulate stomatal closure and other molecules including hydrogen peroxide (H2O2), hydrogen sulfide (H2S), antioxidants, proline, glycine betaine, calcium (Ca2+) and heat shock protein (HSP). Exploring the crosstalk between ABA and NO with other molecules under heat stress will provide us with a comprehensive knowledge of plants mechanism of heat tolerance which could be useful to develop heat stress-resistant varieties.

Crosstalk of plant growth regulators protects photosynthetic performance from arsenic damage by modulating defense systems in rice.

Salicylic acid (SA) is a well-known plant growth regulator, which participates in many physiological processes of plants under normal and stressful conditions. In this study, we investigated the impact of SA supplementation on the components of ascorbate-glutathione cycle and glyoxalase system, photosynthesis and growth of rice (Oryza sativa) plants subjected to arsenic (As) stress. Plants grown with As exhibited enhanced As uptake, increased oxidative stress, and photosynthesis and growth inhibition. Application of SA promoted photosynthesis and growth in plants with or without As stress by improving plant defense systems and reducing oxidative stress through interaction with ethylene and nitric oxide (NO). SA acted as an ethylene antagonist, reducing stress ethylene formation under As stress, while NO formation was induced. This resulted in coordinated control over the antioxidant defense systems and enhanced As tolerance, protecting photosynthesis and growth from As-induced damage. The study showed that positive responses of SA in promoting photosynthesis and growth under As stress were the result of its interplay with ethylene and NO, enhanced capacity of defense systems to reduce oxidative stress. The crosstalk of SA with ethylene and NO will be useful in augmenting the performance of rice plants under As stress.

Treatment of nitric oxide supplemented with nitrogen and sulfur regulates photosynthetic performance and stomatal behavior in mustard under salt stress.

Nitric oxide (NO) is a hormone that connects numerous reactions in plant cells under normal and environmental stress conditions. The application of 100 µM NO as sodium nitroprusside (SNP; NO donor) applied individually or in combination with N or S in different combinations (i.e. 100 mg N or S kg-1 soil applied at the time of sowing [100 N + 100S]0d or with split, 50 mg N or S kg-1 soil at the time of sowing and similar dose at 20 d after sowing [50 N + 50S]0d + [50 N + 50S]20d ) was tested to alleviate salt stress in mustard (Brassica juncea L.). Application of 100 µM NO plus split application of N and S more significantly promoted stomatal behavior, photosynthetic and growth performance in the absence of salt stress and maximally alleviated effects of salt stress through increased N- and S-use efficiency, proline and antioxidant system. The combined application of N and S at the time of sowing was lesser effective in promoting photosynthesis and growth under salt or no salt stress conditions in presence or absence of NO. The study suggests that salt stress effects on the photosynthetic performance are mitigated more efficiently when NO was applied together with the split application of N and S given at two stages, and the photosynthetic activity was promoted under salt stress through increased N and S assimilation and antioxidant system. This strategy may be adopted in agricultural system for overcoming salt stress effects on performance of mustard.

Ethephon mitigates nickel stress by modulating antioxidant system, glyoxalase system and proline metabolism in Indian mustard.

The role of ethylene (through application of ethephon) in the regulation of nickel (Ni) stress tolerance was investigated in this study. Ethephon at concentration of 200 µl l-1 was applied to mustard (Brassica juncea) plants grown without and with 200 mg kg-1 soil Ni to study the increased growth traits, biochemical attributes, photosynthetic efficiency, nutrients content, activities of antioxidants such as superoxide dismutase, ascorbate peroxidase, glutathione reductase, and glutathione peroxidase, glyoxalase systems and enhanced the proline metabolism. In the absence of ethephon, Ni increased oxidative stress with a concomitant decrease in photosynthesis, growth and nutrients content. However, application of ethephon positively increased growth traits, photosynthetic parameters, nutrients content and also elevated the generation of antioxidants enzymes and glyoxalase systems, proline production to combat oxidative stress. Plants water relations and cellular homeostasis were maintained through increased photosynthetic efficiency and proline production. This signifies the role of ethylene in mediating Ni tolerance via regulating proline production and photosynthetic capacity. Ethephon can be used as an exogenous supplement on plants to confer Ni tolerance. The results can be exploited to develop tolerance in plants via gene editing technology encoding enzymes responsible for proline synthesis, antioxidant defence, glyoxalase systems and photosynthetic effectiveness.

Nitric oxide improves S-assimilation and GSH production to prevent inhibitory effects of cadmium stress on photosynthesis in mustard (Brassica juncea L.).

Nitric oxide (NO) is an important gaseous signalling molecule that participates in many developmental and physiological processes, including defense responses against toxic metals in plants. The role of NO in cadmium (Cd)-induced toxic effects on photosynthesis was examined in mustard (Brassica juncea L.) plants. Exposure of plants to 50 µM Cd significantly enhanced oxidative stress (H2O2 content and lipid peroxidation) and impaired plant growth and photosynthesis, chlorophyll fluorescence and reduced chlorophyll content and stomatal conductance. However, the exogenous application of 100 µM sodium nitroprusside (SNP, a donor of NO) reversed the effects of Cd through its stimulation of ROS-scavenging compounds (superoxide dismutase, ascorbate peroxidase, glutathione reductase and reduced glutathione). Exogenous SNP significantly increased plant growth, photosynthesis and chlorophyll content and diminished the accumulation of reactive oxygen species (H2O2) and TBARS content. It also reduced the effects of Cd on thylakoid membrane of the chloroplasts. Application of SNP together with 1.0 mM SO42- showed better responses than SNP alone. The protective effect of NO was achieved through enhanced production of reduced glutathione (GSH). GSH biosynthesis in plants treated with buthionine sulfoximine (BSO), a GSH biosynthetic inhibitor, was not completely inhibited in presence of NO and S, suggesting that NO stimulated S-assimilation and GSH production of Cd exposed plants. This study concludes that NO counteracts Cd toxicity in B. juncea strongly by regulating S-assimilation and GSH production.

Role of Ethylene and Its Cross Talk with Other Signaling Molecules in Plant Responses to Heavy Metal Stress.

Excessive heavy metals (HMs) in agricultural lands cause toxicities to plants, resulting in declines in crop productivity. Recent advances in ethylene biology research have established that ethylene is not only responsible for many important physiological activities in plants but also plays a pivotal role in HM stress tolerance. The manipulation of ethylene in plants to cope with HM stress through various approaches targeting either ethylene biosynthesis or the ethylene signaling pathway has brought promising outcomes. This review covers ethylene production and signal transduction in plant responses to HM stress, cross talk between ethylene and other signaling molecules under adverse HM stress conditions, and approaches to modify ethylene action to improve HM tolerance. From our current understanding about ethylene and its regulatory activities, it is believed that the optimization of endogenous ethylene levels in plants under HM stress would pave the way for developing transgenic crops with improved HM tolerance.

Involvement of ethylene in reversal of salt-inhibited photosynthesis by sulfur in mustard.

Sulfur (S) assimilation results in the synthesis of cysteine (Cys), a common metabolite for the formation of both reduced glutathione (GSH) and ethylene. Thus, ethylene may have regulatory interaction with GSH in the alleviation of salt stress. The involvement of ethylene in the alleviation of salt stress by S application was studied in mustard (Brassica juncea cv. Pusa Jai Kisan). First, the effects of 0, 0.5, 1.0 and 2.0 mM SO4 (2) (-) were studied on photosynthetic and growth parameters to ascertain the S requirement as sufficient-S and excess-S for the plant. In further experiments, the effects of sufficient-S (1 mM SO4 (2) (-) ) and excess-S (2 mM SO4 (2) (-) ) were studied on the alleviation of salt stress-induced by 100 mM NaCl, and ethylene involvement in the alleviation of salt stress by S. Under non-saline condition, excess-S increased ethylene with less content of Cys and GSH and adversely affected photosynthesis and growth. In contrast, excess-S maximally alleviated salt stress due to high demand for S and optimal ethylene formation, which maximally increased GSH and promoted photosynthesis and growth. The involvement of ethylene in S-mediated alleviation of salt stress was further substantiated by the reversal of the effects of excess-S on photosynthesis by aminoethoxyvinylglycine (AVG), ethylene biosynthesis inhibitor. The studies suggest that plants respond differentially to the S availability under non-saline and salt stress and excess-S was more potential in the alleviation of salt stress. Further, ethylene regulates plants' response and excess S-induced alleviation of salt stress and promotion of photosynthesis.

Ethylene production is associated with alleviation of cadmium-induced oxidative stress by sulfur in mustard types differing in ethylene sensitivity.

We studied the response of ethylene-sensitive (Pusa Jai Kisan) and ethylene-insensitive (SS2) mustard (Brassica juncea) cultivars to 0, 0.5, 1.0 and 2.0 mM SO4(2-), and the effect of 1.0 mM SO4(2-) was studied in the amelioration of 50 µM cadmium (Cd). The Cd-induced oxidative stress and Cd accumulation were greater in SS2 than Pusa Jai Kisan, but sulfur (S) application alleviated Cd-induced oxidative stress more prominently in Pusa Jai Kisan by increasing S-metabolism and synthesis of reduced glutathione (GSH) and ethylene production; and promoted photosynthesis and plant dry mass under Cd stress. The ethylene-sensitive cultivar responded more to S treatment under Cd stress and showed increased activity of antioxidant system resulting in increased photosynthesis and growth. Cadmium treatment resulted in rapid increase in ethylene formation which adversely influenced photosynthesis and plant dry mass. However, S and ethephon application to Cd-treated plants lowered ethylene formation to optimal range responsible for maximal GSH synthesis and protection against Cd-induced oxidative stress. The similarity of the effectiveness of 1.0 mM SO4(2-) with 200 µL L(-1) ethylene source as ethephon in alleviation of 50 µM Cd further verifies that differential alleviation of Cd toxicity in the two cultivars by S was dependent on ethylene production. The results suggest that ethylene production determines Cd stress alleviation by S via regulatory interaction with antioxidant metabolism. Thus, ethylene production and sensitivity bear a prominent role in alleviation of Cd stress by S and can be used as a criterion for developing Cd tolerant genotypes.

Appraisal of the Role of Gaseous Signaling Molecules in Thermo-Tolerance Mechanisms in Plants.

A significant threat to the ongoing rise in temperature caused by global warming. Plants have many stress-resistance mechanisms, which is responsible for maintaining plant homeostasis. Abiotic stresses largely increase gaseous molecules' synthesis in plants. The study of gaseous signaling molecules has gained attention in recent years. The role of gaseous molecules, such as nitric oxide (NO), hydrogen sulfide (H2S), carbon dioxide (CO2), carbon monoxide (CO), methane (CH4), and ethylene, in plants under temperature high-temperature stress are discussed in the current review. Recent studies revealed the critical function that gaseous molecules play in controlling plant growth and development and their ability to respond to various abiotic stresses. Here, we provide a thorough overview of current advancements that prevent heat stress-related plant damage via gaseous molecules. We also explored and discussed the interaction of gaseous molecules. In addition, we provided an overview of the role played by gaseous molecules in high-temperature stress responses, along with a discussion of the knowledge gaps and how this may affect the development of high-temperature-resistant plant species.

Hydrogen sulfide and ethylene regulate sulfur-mediated stomatal and photosynthetic responses and heat stress acclimation in rice.

The gaseous signaling molecules, ethylene (ET) and hydrogen sulfide (H2S) are well known for their ability to mitigate abiotic stress, but how they interact with mineral nutrients under heat stress is unclear. We have studied the involvement of ET and H2S in adaptation of heat stress on the availability of sulfur (S) levels in rice (Oryza sativa L.). Heat stress (40 °C) negatively impacted growth and photosynthetic-sulfur use efficiency (p-SUE), with accumulation of reactive oxygen species (ROS) in six rice cultivars, namely PS 2511, Birupa, Nidhi, PB 1509, PB 1728, and Panvel. Supplementation of S at 2.0 mM SO42- in the form of MgSO4, improved growth and photosynthetic attributes more than 1.0 mM SO42- under control (28 °C), and mitigated heat stress effects more prominently in PS 2511 (heat-tolerant) than in PB 1509 (heat-sensitive) cultivar. The higher heat stress mitigation potential of 2.0 mM SO42- in heat-tolerant cultivar was correlated with higher S-assimilation, activity of antioxidant enzymes, stomatal (stomatal conductance) and non-stomatal limitations, activity of carbonic anhydrase and Rubisco, and mesophyll conductance. The use of norbornadiene (NBD) and hypotaurine (HT), ET and H2S inhibitors, respectively, resulted in the lowest values for photosynthetic efficiency, stomatal and non-stomatal factors, implying the mediation of ET and H2S in heat stress acclimation. The connectivity of ET and H2S with S-assimilation through a common metabolite cysteine (Cys) improved heat stress adaptation in which H2S acted downstream to ET-mediated responses. Thus, the better adaptability of rice plants to heat stress may be obtained through modulation of ET and H2S via S.

Multifaceted roles of silicon nano particles in heavy metals-stressed plants.

Heavy metal (HM) contamination has emerged as one of the most damaging abiotic stress factors due to their prominent release into the environment through industrialization and urbanization worldwide. The increase in HMs concentration in soil and the environment has invited attention of researchers/environmentalists to minimize its' impact by practicing different techniques such as application of phytohormones, gaseous molecules, metalloids, and essential nutrients etc. Silicon (Si) although not considered as the essential nutrient, has received more attention in the last few decades due to its involvement in the amelioration of wide range of abiotic stress factors. Silicon is the second most abundant element after oxygen on earth, but is relatively lesser available for plants as it is taken up in the form of mono-silicic acid, Si(OH)4. The scattered information on the influence of Si on plant development and abiotic stress adaptation has been published. Moreover, the use of nanoparticles for maintenance of plant functions under limited environmental conditions has gained momentum. The current review, therefore, summarizes the updated information on Si nanoparticles (SiNPs) synthesis, characterization, uptake and transport mechanism, and their effect on plant growth and development, physiological and biochemical processes and molecular mechanisms. The regulatory connect between SiNPs and phytohormones signaling in counteracting the negative impacts of HMs stress has also been discussed.

The Ethylene Biosynthetic Enzymes, 1-Aminocyclopropane-1-Carboxylate (ACC) Synthase (ACS) and ACC Oxidase (ACO): The Less Explored Players in Abiotic Stress Tolerance.

Ethylene is an essential plant hormone, critical in various physiological processes. These processes include seed germination, leaf senescence, fruit ripening, and the plant's response to environmental stressors. Ethylene biosynthesis is tightly regulated by two key enzymes, namely 1-aminocyclopropane-1-carboxylate synthase (ACS) and 1-aminocyclopropane-1-carboxylate oxidase (ACO). Initially, the prevailing hypothesis suggested that ACS is the limiting factor in the ethylene biosynthesis pathway. Nevertheless, accumulating evidence from various studies has demonstrated that ACO, under specific circumstances, acts as the rate-limiting enzyme in ethylene production. Under normal developmental processes, ACS and ACO collaborate to maintain balanced ethylene production, ensuring proper plant growth and physiology. However, under abiotic stress conditions, such as drought, salinity, extreme temperatures, or pathogen attack, the regulation of ethylene biosynthesis becomes critical for plants' survival. This review highlights the structural characteristics and examines the transcriptional, post-transcriptional, and post-translational regulation of ACS and ACO and their role under abiotic stress conditions. Reviews on the role of ethylene signaling in abiotic stress adaptation are available. However, a review delineating the role of ACS and ACO in abiotic stress acclimation is unavailable. Exploring how particular ACS and ACO isoforms contribute to a specific plant's response to various abiotic stresses and understanding how they are regulated can guide the development of focused strategies. These strategies aim to enhance a plant's ability to cope with environmental challenges more effectively.

Gamma-aminobutyric acid interactions with phytohormones and its role in modulating abiotic and biotic stress in plants.

Gamma-aminobutyric acid (GABA), a ubiquitous non-protein 4-carbon amino acid present in both prokaryotic and eukaryotic organisms. It is conventionally recognized as a neurotransmitter in mammals and plays a crucial role in plants. The context of this review centers on the impact of GABA in mitigating abiotic stresses induced by climate change, such as drought, salinity, heat, and heavy metal exposure. Beyond its neurotransmitter role, GABA emerges as a key player in diverse metabolic processes, safeguarding plants against multifaceted abiotic as well as biotic challenges. This comprehensive exploration delves into the GABA biosynthetic pathway, its transport mechanisms, and its intricate interplay with various abiotic stresses. The discussion extends to the nuanced relationship between GABA and phytohormones during abiotic stress acclimation, offering insights into the strategic development of mitigation strategies against these stresses. The delineation of GABA's crosstalk with phytohormones underscores its pivotal role in formulating crucial strategies for abiotic stress alleviation in plants.

Relative effects of melatonin and hydrogen sulfide treatments in mitigating salt damage in wheat.

Soil salinity poses a significant threat to agricultural productivity, impacting the growth and yield of wheat (Triticum aestivum L.) plants. This study investigates the potential of melatonin (MT; 100 µM) and hydrogen sulfide (H2S; 200 µM sodium hydrosulfide, NaHS) to confer the tolerance of wheat plants to 100 mM NaCl. Salinity stress induced the outburst of reactive oxygen species (ROS) resulting in damage to the chloroplast structure, growth, photosynthesis, and yield. Application of either MT or NaHS augmented the activity of antioxidant enzymes, superoxide dismutase, ascorbate peroxidase, glutathione reductase, and reduced glutathione (GSH) levels, upregulated the expression of Na+ transport genes (SOS1, SOS2, SOS3, NHX1), resulting in mitigation of salinity stress. Thus, improved stomatal behavior, gas-exchange parameters, and maintenance of chloroplast structure resulted in enhanced activity of the Calvin cycle enzymes and overall enhancement of growth, photosynthetic, and yield performance of plants under salinity stress. The use of DL-propargylglycine (PAG, an inhibitor of hydrogen sulfide biosynthesis) and p-chlorophenyl alanine (p-CPA, an inhibitor of melatonin biosynthesis) to plants under salt stress showed the comparative necessity of MT and H2S in mitigation of salinity stress. In the presence of PAG, more pronounced detrimental effects were observed than in the presence of p-CPA, emphasizing that MT was involved in mitigating salinity through various potential pathways, one of which was through H2S.

Nitric Oxide and Proline Modulate Redox Homeostasis and Photosynthetic Metabolism in Wheat Plants under High Temperature Stress Acclimation.

The effects of exogenously-sourced NO (nitric oxide, as 100 µM SNP) and proline (50 mM) in the protection of the photosynthetic performance of wheat (Triticum aestivum L.) plants against heat stress were investigated. The study focused on the mechanisms of proline accumulation, activity, gene expression of antioxidant enzymes, and NO generation. Plants were exposed to a temperature of 40 °C for 6 h per day over 15 days, then allowed to recover at 28 °C. Heat-stressed plants showed increased oxidative stress, with higher levels of H2O2 and TBARS (thiobarbituric acid reactive substances) and increased proline accumulation, ACS activity, ethylene evolution, and NO generation, which in turn leads to increased accumulation of antioxidant enzymes and reduced photosynthetic attributes. In the tested wheat cultivar, the exogenous application of SNP and proline under heat stress improved the photosynthesis and reduced oxidative stress by enhancing the enzymatic antioxidant defense system. Potentially, the promoter AOX (alternative oxidase) played a role in maintaining redox homeostasis by lowering H2O2 and TBARS levels. The genes for GR antioxidant and photosystem II core protein encoding psbA and psbB were highly up-regulated in nitric oxide and proline treated heat-stressed plants, indicating that ethylene positively impacted photosynthesis under high temperature stress. Moreover, nitric oxide supplementation under high temperature stress optimized ethylene levels to regulate the assimilation and metabolism of proline and the antioxidant system, lowering the adverse effects. The study showed that nitric oxide and proline increased high temperature stress tolerance in wheat by increasing the osmolytes accumulation and the antioxidant system, resulting in enhanced photosynthesis.

Review on nitric oxide at the forefront of rapid systemic signaling in mitigation of salinity stress in plants: Crosstalk with calcium and hydrogen peroxide.

Soil salinity is a global issue that limits plant growth in agricultural fields and contributes to food crisis. Salt stressors impede plant's ionic, osmotic, and oxidative balance, as well as a variety of physiological functions. Exposure to salinity stress manifest considerable ROS clustering, entailing modification in performance of various organelles. To deal with salinity, plants use a variety of coping strategies, such as osmoregulation, ion-homeostasis, increased antioxidant synthesis, and so on. Nitric oxide (NO) is a pivotal signalling molecule that helps facilitate salt stress-induced physiological plant responses. A variety of evidences point to NO being produced under similar stress conditions and with similar kinetics as hydrogen peroxide (H2O2). The interplay between H2O2 and NO has important functional implications for modulating plant transduction processes. Besides, NO and calcium (Ca2+)-dependent pathways also have some connection in salt stress response mechanisms. Extensive crosstalk between NO and Ca2+ signalling pathways is investigated, and it suggests that almost every type of Ca2+ channel is under the tight control of NO, and NO acts as a Ca2+ mobilising compound and aids in signal reliance. The review provides insights into understanding recent advances regarding NO's, Ca2+ and H2O2 role in salt stress reduction with entwine signaling mechanisms.