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.

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.

Melatonin Reverses High-Temperature-Stress-Inhibited Photosynthesis in the Presence of Excess Sulfur by Modulating Ethylene Sensitivity in Mustard.

Melatonin is a pleiotropic, nontoxic, regulatory biomolecule with various functions in abiotic stress tolerance. It reverses the adverse effect of heat stress on photosynthesis in plants and helps with sulfur (S) assimilation. Our research objective aimed to find the influence of melatonin, along with excess sulfur (2 mM SO42-), in reversing heat stress's impacts on the photosynthetic ability of the mustard (Brassica juncea L.) cultivar SS2, a cultivar with low ATP-sulfurylase activity and a low sulfate transport index (STI). Further, we aimed to substantiate that the effect was a result of ethylene modulation. Melatonin in the presence of excess-S (S) increased S-assimilation and the STI by increasing the ATP-sulfurylase (ATP-S) and serine acetyltransferase (SAT) activity of SS2, and it enhanced the content of cysteine (Cys) and methionine (Met). Under heat stress, melatonin increased S-assimilation and diverted Cys towards the synthesis of more reduced glutathione (GSH), utilizing excess-S at the expense of less methionine and ethylene and resulting in plants' reduced sensitivity to stress ethylene. The treatment with melatonin plus excess-S increased antioxidant enzyme activity, photosynthetic-S use efficiency (p-SUE), Rubisco activity, photosynthesis, and growth under heat stress. Further, plants receiving melatonin and excess-S in the presence of norbornadiene (NBD; an ethylene action inhibitor) under heat stress showed an inhibited STI and lower photosynthesis and growth. This suggested that ethylene was involved in the melatonin-mediated heat stress reversal effects on photosynthesis in plants. The interaction mechanism between melatonin and ethylene is still elusive. This study provides avenues to explore the melatonin-ethylene-S interaction for heat stress tolerance in plants.

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.

Melatonin influences methyl jasmonate-induced protection of photosynthetic activity in wheat plants against heat stress by regulating ethylene-synthesis genes and antioxidant metabolism.

Melatonin (MT) and methyl jasmonate (MeJA) play important roles in the adaptation of plants to different stress factors by modulating stress tolerance mechanisms. The present study reports the involvement of MT (100 µM) in MeJA (10 µM)-induced photosynthetic performance and heat stress acclimation through regulation of the antioxidant metabolism and ethylene production in wheat (Triticum aestivum L.) plants. Plants exposed to 40 °C for 6 h per day for 15 days and allowed to retrieve at 28 °C showed enhanced oxidative stress and antioxidant metabolism, increased 1-aminocyclopropane-1-carboxylic acid (ACC) synthase (ACS) activity and ethylene production, and decreased photosynthetic performance. In contrast, the exogenously applied MT and MeJA reduced oxidative stress through improved S-assimilation (+ 73.6% S content), antioxidant defense system (+ 70.9% SOD, + 115.8% APX and + 104.2% GR, and + 49.5% GSH), optimized ethylene level to 58.4% resulting in improved photosynthesis by 75%. The use of p-chlorophenyl alanine, a MT biosynthesis inhibitor along with MeJA in the presence of heat stress reduced the photosynthetic performance, ATP-S activity and GSH content, substantiated the requirement of MT in the MeJA-induced photosynthetic response of plants under heat stress. These findings suggest that MeJA evoked the plant's ability to withstand heat stress by regulating the S-assimilation, antioxidant defense system, and ethylene production, and improving photosynthetic performance was dependent on MT.

Sulfur supplementation enhances nitric oxide efficacy in reversal of chromium-inhibited Calvin cycle enzymes, photosynthetic activity, and carbohydrate metabolism in wheat.

The present study demonstrated that exogenously-sourced nitric oxide (as SNP, sodium nitroprusside; NO donor) and sulfur (S) protected photosynthesis against chromium (Cr) stress in wheat (Triticum aestivum L. cv. HD 2851). Plants grown with 100 µM Cr exhibited higher reactive oxygen species (ROS) production, resulting in photosynthetic damage. The individual application of 50 µM NO increased carbohydrate metabolism as well as photosynthetic parameters, antioxidant system with higher transcriptional gene levels that encode the key enzymes for the Calvin cycle under Cr stress. These effects were more prominent when NO was applied with 1.0 mM SO42-. An increase in the reduced glutathione (GSH) content obtained with NO was further enhanced by S and resulted in higher protection against Cr stress. The protective effect of NO with S against Cr toxicity on photosynthesis was reversed when buthionine sulfoximine (BSO; GSH biosynthetic inhibitor) was used. Application of BSO reversed the impact of NO plus S on photosynthesis under Cr stress, verifying that the ameliorating effect of NO was through S-assimilation and via GSH production. Thus, the availability of S to NO application can help reduce Cr toxicity and protect photosynthetic activity and expression of the Calvin cycle enzymes in leaves through the GSH involvement.

Nitric Oxide and Salicylic Acid Regulate Glutathione and Ethylene Production to Enhance Heat Stress Acclimation in Wheat Involving Sulfur Assimilation.

Phytohormones have a role in stress adaptation. The major mechanism underlying the role of exogenously-sourced nitric oxide (NO; as sodium nitroprusside, SNP: 50.0 µM) and salicylic acid (SA; 0.5 mM) in the presence of 2.0 mM SO4-2 was assessed in heat stress (HS; 40 °C for 6 h daily for 15 days) tolerance in wheat (Triticum aestivum L. cv. HD-3226). The cultivar HD-3226 possessed high photosynthetic sulfur use efficiency (p-SUE) among the six cultivars screened. Plants grown under HS exhibited an increased content of reactive oxygen species (ROS; including superoxide radical and hydrogen peroxide) and extent of lipid peroxidation with a consequent reduction in photosynthesis and growth. However, both NO and SA were found to be protective against HS via enhanced S assimilation. Their application reduced oxidative stress and increased the activity of antioxidant enzymes. NO or SA supplementation along with S under HS recovered the losses and improved photosynthesis and growth. The use of SA inhibitor (2-aminoindane-2-phosphonic acid; AIP) and NO scavenger (cPTIO) confirmed that the mitigating effects of SA and NO involved induction in S assimilation.

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.

Ethylene Signaling under Stressful Environments: Analyzing Collaborative Knowledge.

Ethylene is a gaseous plant growth hormone that regulates various plant developmental processes, ranging from seed germination to senescence. The mechanisms underlying ethylene biosynthesis and signaling involve multistep mechanisms representing different control levels to regulate its production and response. Ethylene is an established phytohormone that displays various signaling processes under environmental stress in plants. Such environmental stresses trigger ethylene biosynthesis/action, which influences the growth and development of plants and opens new windows for future crop improvement. This review summarizes the current understanding of how environmental stress influences plants' ethylene biosynthesis, signaling, and response. The review focuses on (a) ethylene biosynthesis and signaling in plants, (b) the influence of environmental stress on ethylene biosynthesis, (c) regulation of ethylene signaling for stress acclimation, (d) potential mechanisms underlying the ethylene-mediated stress tolerance in plants, and (e) summarizing ethylene formation under stress and its mechanism of action.

Hydrogen Sulfide, Ethylene, and Nitric Oxide Regulate Redox Homeostasis and Protect Photosynthetic Metabolism under High Temperature Stress in Rice Plants.

Rising temperatures worldwide due to global climate change are a major scientific issue at present. The present study reports the effects of gaseous signaling molecules, ethylene (200 µL L-1; 2-chloroethylphosphonic acid; ethephon, Eth), nitric oxide (NO; 100 µM sodium nitroprusside; SNP), and hydrogen sulfide (H2S; 200 µM sodium hydrosulfide, NaHS) in high temperature stress (HS) tolerance, and whether or not H2S contributes to ethylene or NO-induced thermo-tolerance and photosynthetic protection in rice (Oryza sativa L.) cultivars, i.e., Taipei-309, and Rasi. Plants exposed to an HS of 40 °C for six h per day for 15 days caused a reduction in rice biomass, associated with decreased photosynthesis and leaf water status. High temperature stress increased oxidative stress by increasing the content of hydrogen peroxide (H2O2) and thiobarbituric acid reactive substance (TBARS) in rice leaves. These signaling molecules increased biomass, leaf water status, osmolytes, antioxidants, and photosynthesis of plants under non-stress and high temperature stress. However, the effect was more conspicuous with ethylene than NO and H2S. The application of H2S scavenger hypotaurine (HT) reversed the effect of ethylene or NO on photosynthesis under HS. This supports the findings that the ameliorating effects of Eth or SNP involved H2S. Thus, the presence of H2S with ethylene or NO can enhance thermo-tolerance while also protecting plant photosynthesis.

Ethylene Suppresses Abscisic Acid, Modulates Antioxidant System to Counteract Arsenic-Inhibited Photosynthetic Performance in the Presence of Selenium in Mustard.

Arsenic (As) stress provokes various toxic effects in plants that disturbs its photosynthetic potential and hampers growth. Ethylene and selenium (Se) have shown regulatory interaction in plants for metal tolerance; however, their synergism in As tolerance through modification of the antioxidant enzymes and hormone biosynthesis needs further elaboration. With this in view, we investigated the impact of ethylene and Se in the protection of photosynthetic performance against As stress in mustard (Brassica juncea L.). Supplementation with ethephon (2-chloroethylphosphonic acid; ethylene source) and/or Se allayed the negative impact of As-induced toxicity by limiting As content in leaves, enhancing the antioxidant defense system, and decreasing the accumulation of abscisic acid (ABA). Ethylene plus Se more prominently regulated stomatal behavior, improved photosynthetic capacity, and mitigated As-induced effects. Ethephon in the presence of Se decreased stress ethylene formation and ABA accumulation under As stress, resulting in improved photosynthesis and growth through enhanced reduced glutathione (GSH) synthesis, which in turn reduced the oxidative stress. In both As-stressed and non-stressed plants treated with ethylene action inhibitor, norbornadiene, resulted in increased ABA and oxidative stress with reduced photosynthetic activity by downregulating expression of ascorbate peroxidase and glutathione reductase, suggesting the involvement of ethylene in the reversal of As-induced toxicity. These findings suggest that ethephon and Se induce regulatory interaction between ethylene, ABA accumulation, and GSH metabolism through regulating the activity and expression of antioxidant enzymes. Thus, in an economically important crop (mustard), the severity of As stress could be reduced through the supplementation of both ethylene and Se that coordinate for maximum stress alleviation.

The Functional Interplay between Ethylene, Hydrogen Sulfide, and Sulfur in Plant Heat Stress Tolerance.

Plants encounter several abiotic stresses, among which heat stress is gaining paramount attention because of the changing climatic conditions. Severe heat stress conspicuously reduces crop productivity through changes in metabolic processes and in growth and development. Ethylene and hydrogen sulfide (H2S) are signaling molecules involved in defense against heat stress through modulation of biomolecule synthesis, the antioxidant system, and post-translational modifications. Other compounds containing the essential mineral nutrient sulfur (S) also play pivotal roles in these defense mechanisms. As biosynthesis of ethylene and H2S is connected to the S-assimilation pathway, it is logical to consider the existence of a functional interplay between ethylene, H2S, and S in relation to heat stress tolerance. The present review focuses on the crosstalk between ethylene, H2S, and S to highlight their joint involvement in heat stress tolerance.

Ethylene-nitrogen synergism induces tolerance to copper stress by modulating antioxidant system and nitrogen metabolism and improves photosynthetic capacity in mustard.

This study aimed to test the efficiency of ethylene (Eth; 200 µL L-1 ethephon) in presence or absence of nitrogen (N; 80 mg N kg-1 soil) in protecting photosynthetic apparatus from copper (Cu; 100 mg Cu kg-1 soil) stress in mustard (Brassica juncea L.) and to elucidate the physio-biochemical modulation for Eth plus N-induced Cu tolerance. Elevated Cu-accrued reductions in photosynthesis and growth were accompanied by significantly higher Cu accumulation in leaves and oxidative stress with reduced assimilation of N and sulfur (S). Ethylene in coordination with N considerably reduced Cu accumulation, lowered lipid peroxidation, lignin accumulation, and contents of reactive oxygen species (hydrogen peroxide, H2O2, and superoxide anion, O2•-), and mitigated the negative effect of Cu on N and S assimilation, accumulation of non-protein thiols and phytochelatins, enzymatic, and non-enzymatic antioxidants (activity of ascorbate peroxidase, APX, and glutathione reductase, GR; content of reduced glutathione, GSH, and ascorbate, AsA), cell viability, photosynthesis, and growth. Overall, the effect of ethylene-nitrogen synergism was evident on prominently mitigating Cu stress and protecting photosynthesis. The approach of supplementing ethylene with N may be used as a potential tool to restrain Cu stress, and protect photosynthesis and growth of mustard plants.

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.

Ethylene Supplementation Combined with Split Application of Nitrogen and Sulfur Protects Salt-Inhibited Photosynthesis through Optimization of Proline Metabolism and Antioxidant System in Mustard (Brassica juncea L.).

In the present study, the potential of ethylene as ethephon (an ethylene source) was investigated individually and in combination with split doses of nitrogen (N) and sulfur (S) soil treatments for removal of the damaging effects of salt stress (100 mM NaCl) in mustard (Brassica juncea L.). Plants were grown with 50 mg N plus 50 mg S kg-1 soil at sowing time and an equivalent dose at 20 days after sowing [N50 + S50]0d and 20d. Ethephon at 200 µL L‒1 was applied to combined split doses of N and S with or without NaCl. Plants subjected to NaCl showed a decrease in growth and photosynthetic characteristics as well as N and S assimilation, whereas proline metabolism and antioxidants increased. The application of ethephon to plants grown with split N and S doses significantly enhanced photosynthetic efficiency by increasing the assimilation of N and S, improving the concentration of proline and induction of the antioxidant system with or without NaCl. The regulation of ethylene and/or split forms of N and S application may be potential tools for not just overcoming salt stress effects in this species and in related Brassicaceae but also enhancing their photosynthesis and growth potential through increased nutrient assimilation.

Ethylene reduces glucose sensitivity and reverses photosynthetic repression through optimization of glutathione production in salt-stressed wheat (Triticum aestivum L.).

Ethylene plays a crucial role throughout the life cycle of plants under optimal and stressful environments. The present study reports the involvement of exogenously sourced ethylene (as ethephon; 2-chloroethyl phosphonic acid) in the protection of the photosynthetic activity from glucose (Glu) sensitivity through its influence on the antioxidant system for adaptation of wheat (Triticum aestivum L.) plants under salt stress. Ten-day-old plants were subjected to control and 100 mM NaCl and treated with 200 µl L-1 ethephon on foliage at 20 days after seed sowing individually or in combination with 6% Glu. Plants receiving ethylene exhibited higher growth and photosynthesis through reduced Glu sensitivity in the presence of salt stress. Moreover, ethylene-induced reduced glutathione (GSH) production resulted in increased psbA and psbB expression to protect PSII activity and photosynthesis under salt stress. The use of buthionine sulfoximine (BSO), GSH biosynthesis inhibitor, substantiated the involvement of ethylene-induced GSH in the reversal of Glu-mediated photosynthetic repression in salt-stressed plants. It was suggested that ethylene increased the utilization of Glu under salt stress through its influence on photosynthetic potential and sink strength and reduced the Glu-mediated repression of photosynthesis.

Control of Elevated Ion Accumulation, Oxidative Stress, and Lipid Peroxidation with Salicylic Acid-Induced Accumulation of Glycine Betaine in Salinity-Exposed Vigna radiata L.

The identification of the sustainable approaches is required for the minimization of adverse impact of worldwide increasing soil salinity on plant growth, development, and productivity. This study investigated the protective role and major mechanism underlying salicylic acid (SA; 0.1, 0.5, or 1.0 mM)-induced glycine betaine (GB)-mediated tolerance to salinity (50 mM NaCl) in mungbean (Vigna radiata L. cultivar Punt Mung). The supply of 0.5 mM SA maximally increased the accumulation of GB (>40%) with respect to the control. This was further corroborated with the increase in water potential, antioxidant system (reduced glutathione (GSH), GSH/GSSG redox state, and glutathione reductase (GR) activity) and decreased Na+ and Cl- accumulation, Na+/K+ ratio, oxidative stress, and lipid peroxidation. This was also associated with the increased photosynthesis (14-18%) and growth (7-12%) parameters. Overall, SA-induced accumulation of GB protected photosynthesis and growth against 50 mM NaCl-accrued impacts in V. radiata through minimizing the accumulation of Na+ and Cl- ions, oxidative stress, and maintaining high GSH level that led to reduced cellular redox environment.

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.

Nitric Oxide Enhances Photosynthetic Nitrogen and Sulfur-Use Efficiency and Activity of Ascorbate-Glutathione Cycle to Reduce High Temperature Stress-Induced Oxidative Stress in Rice (Oryza sativa L.) Plants.

The effects of nitric oxide (NO) as 100 µM sodium nitroprusside (SNP, NO donor) on photosynthetic-nitrogen use efficiency (NUE), photosynthetic-sulfur use efficiency (SUE), photosynthesis, growth and agronomic traits of rice (Oryza sativa L.) cultivars, Taipie-309 (high photosynthetic-N and SUE) and Rasi (low photosynthetic-N and SUE) were investigated under high temperature stress (40 °C for 6 h). Plants exposed to high temperature stress caused significant reduction in photosynthetic activity, use efficiency of N and S, and increment in H2O2 and thiobarbituric acid reactive substance (TBARS) content. The drastic effects of high temperature stress were more pronounced in cultivar Rasi than Taipie-309. However, foliar spray of SNP decreased the high temperature induced H2O2 and TBARS content and increased accumulation of proline and activity of ascorbate-glutathione cycle that collectively improved tolerance to high temperature stress more effectively in Taipie-309. Exogenously applied SNP alleviated the high temperature induced decrease in photosynthesis through maintaining higher photosynthetic-NUE and photosynthetic-SUE, activity of ribulose 1,5 bisphosphate carboxylase/oxygenase (Rubisco), and synthesis of reduced glutathione (GSH). The use of 2-4-carboxyphenyl-4,4,5,5-tetramethylimidazoline-1-oxy-3-oxide (cPTIO, NO scavenger) substantiated the study that in the absence of NO oxidative stress increased, while NO increased photosynthetic-NUE and photosynthetic-SUE, net photosynthesis and plant dry mass. Taken together, the present investigation reveals that NO increased heat stress tolerance and minimized high temperature stress adversaries more effectively in cultivar Taipie-309 than Rasi by enhancing photosynthetic-NUE and SUE and strengthening the antioxidant defense system.