Search

Plants face a range of environmental stresses, such as heat and drought, that significantly reduce their growth, development, and yield. Plants have developed complex signaling networks to regulate physiological processes and improve their ability to withstand stress. The key regulators of plant stress responses include polyamines (PAs) and gaseous signaling molecules (GSM), such as hydrogen sulfide (H 2 S), nitric oxide (NO), methane (CH 4 ), carbon monoxide (CO), carbon dioxide (CO 2 ), and ethylene (ET). The functions of PAs and GSM in stress perception, signal transduction, and stress-responsive pathways have been explored. However, there is a lack of detailed, updated information on the interaction of PAs and GSM in the adaptation of drought and heat stress. This review explores the interaction between PAs and GSM for the adaptation to drought and heat stress. It explores their synergistic effects in mitigating the negative impacts of drought and heat stress on plant growth, development, and productivity. Moreover, a comprehensive analysis of physiological, biochemical, and molecular approaches demonstrates that their interaction activates key stress-responsive pathways, enhances antioxidant systems, and modulates gene expression. These combined effects contribute to improved drought and heat tolerance in plants. The information presented in the review provides valuable insights into plant stress resilience strategies and suggests potential measures for developing climate-resilient crops to address the increasing environmental challenges.

Plants are frequently exposed to high salinity, negatively affecting their development and productivity. This review examined the complex roles of membrane transporters (MTs) and phytohormones in mediating salt stress. MTs are crucial in capturing sodium ions (Na + ) and maintaining a delicate balance between sodium (Na + ) and potassium (K + ), essential for supporting cellular homeostasis and enhancing overall plant health. These MTs were instrumental in regulating ion balance and promoting the absorption and segregation of vital nutrients, thereby enhancing salt stress tolerance. Various plant hormones, including abscisic acid, auxin, ethylene, cytokinin, and gibberellins, along with gaseous growth regulators such as nitric oxide and hydrogen sulfide, collaborate to regulate and synchronize numerous aspects of plant growth, development, and stress responses to environmental factors. These transporters and other phytohormones, including brassinosteroids, melatonin, and salicylic acid, also collaborated to initiate adaptation processes, such as controlling osmotic pressure, removing ions, and initiating stress signaling pathways. This study consolidated the advancements in understanding the molecular and physiological processes contributing to plant salt tolerance, emphasizing the intricate relationships between MTs and phytohormones. The aim was to elucidate these interactions to promote further research and develop strategies for enhancing plant salt tolerance., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could, that could have appeared to influence the work reported in this article., (Copyright © 2025 Elsevier Masson SAS. All rights reserved.)

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.

Salt stress is an alarming abiotic stress that reduces mustard growth and yield. To attenuate salt toxicity effects, plant growth-promoting rhizobacteria (PGPR) offers a sustainable approach. Among the various PGPR, Pseudomonas fluorescens (P. fluorescens NAIMCC-B-00340) was chosen for its salt tolerance (at 100 mM NaCl) and for exhibiting various growth-promoting activities. Notably, P. fluorescens can produce auxin, which plays a role in melatonin (MT) synthesis. Melatonin is a pleiotropic molecule that acts as an antioxidant to scavenge reactive oxygen species (ROS), resulting in stress reduction. Owing to the individual role of PGPR and MT in salt tolerance, and their casual nexus, their domino effect was investigated in Indian mustard under salt stress. The synergistic action of P. fluorescens and MT under salt stress conditions was found to enhance the activity of antioxidative enzymes and proline content as well as  promote the production of secondary metabolites. This led to reduced oxidative stress following effective ROS scavenging, maintained photosynthesis, and improved growth. In mustard plants treated with MT and P. fluorescens under salt stress, eight flavonoids showed significant increase. Kaempferol and cyanidin showed the highest concentrations and are reported to act as antioxidants with protective functions under stress. Thus, we can anticipate that strategies involved in their enhancement could provide a better adaptive solution to salt toxicity in mustard plants. In conclusion, the combination of P. fluorescens and MT affected antioxidant metabolism and flavonoid profile that could be used to mitigate salt-induced stress and bolster plant resilience., (© 2023 Scandinavian Plant Physiology Society.)

This study presents an exploration of the efficacy of brassinosteroids (BRs) and ethylene in mediating heat stress tolerance in rice (Oryza sativa). Heat is one of the major abiotic factors that prominently deteriorates rice production by influencing photosynthetic efficiency, source‒sink capacity, and growth traits. The application of BR (0.5 mM) and ethylene (200 μl l -1 ) either individually and/or in combination was found to alleviate heat stress-induced toxicity by significantly improving photosynthesis, source‒sink capacity and defense systems; additionally, it reduced the levels of oxidative stress markers and ethylene formation. The study revealed the positive influence of BR in promoting plant growth responses under heat stress through its interplay with ethylene biosynthesis and enhanced plant defense systems. Interestingly, treatment with the ethylene biosynthesis inhibitor aminoethoxyvinylglycine (AVG) substantiated that BR application to heat-stressed rice plants enhanced ethylene-dependent pathways to counteract the underlying adversities. Thus, BR action was found to be mediated by ethylene to promote heat tolerance in rice. The present study sheds light on the potential tolerance mechanisms which can ensure rice sustainability under heat stress conditions., Competing Interests: Declaration of competing interest Authors declare that they have no conflict of interest., (Copyright © 2023 Elsevier GmbH. All rights reserved.)

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 SO 4 2- ), 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 plant growth regulator, jasmonic acid (JA) has emerged as important molecule and involved in key processes of plants. In this study, we investigated the role of methyl jasmonate (MeJA) in achieving tolerance mechanisms against arsenic (As) stress in rice (Oryza sativa). Arsenic toxicity is a major global concern that significantly deteriorate rice production. The application of MeJA (20 μM) and ethylene (150 μL L -1 ) both individually and/or in combination were found significant in protecting against As-induced toxicity in rice, and significantly improved defense systems. The study shown that the positive influence of MeJA in promoting carbohydrate metabolism, photosynthesis and growth under As stress were the result of its interplay with ethylene biosynthesis and reduced oxidative stress-mediated cellular injuries and cell deaths. Interestingly, the use of JA biosynthesis inhibitor, neomycin (Neo) and ethylene biosynthesis inhibitor, aminoethoxyvinylglycine (AVG) overturned the effects of MeJA and ethylene on plant growth under As stress. From the pooled data, it may also be concluded that Neo treatment to MeJA- treated rice plants restricted JA-mediated responses, implying that application of MeJA modulated ethylene- dependent pathways in response to As stress. Thus, the action of MeJA in As tolerance is found to be mediated by ethylene. The study will shed light on the mechanisms that could be used to ensure the sustainability of rice plants under As stress., Competing Interests: Declaration of competing interest All authors have read and agreed to the published version of the manuscript. Authors have no conflicts., (Copyright © 2023 Elsevier Masson SAS. All rights reserved.)

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 (H 2 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., (© 2023 Scandinavian Plant Physiology Society.)

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 SO 4 2- . 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., (© 2023. The Author(s).)

The severity of salt stress is alarming for crop growth and production and it threatens food security. Strategies employed for the reduction in stress are not always eco-friendly or sustainable. Plant-growth-promoting rhizobacteria (PGPR) could provide an alternative sustainable stress reduction strategy owning to its role in various metabolic processes. In this study, we have used two strains of PGPR, Pseudomonas fluorescens (NAIMCC-B-00340) and Azotobacter chroococcum Beijerinck 1901 (MCC 2351), either singly or in combination, and studied their effect in the amelioration of salt toxicity in mustard cultivar Pusa Jagannath via its influence on plants' antioxidants' metabolism, photosynthesis and growth. Individually, the impact of Pseudomonas fluorescens was better in reducing stress ethylene, oxidative stress, photosynthesis and growth but maximal alleviation was observed with their combined application. MDA and H 2 O 2 content as indicator of oxidative stress decreased by 27.86% and 45.18% and osmolytes content (proline and glycine-betaine) increased by 38.8% and 26.3%, respectively, while antioxidative enzymes (SOD, CAT, APX and GR) increased by 58.40, 25.65, 81.081 and 55.914%, respectively, over salt-treated plants through the application of Pseudomonas fluorescens . The combined application maximally resulted in more cell viability and less damage to the leaf with lesser superoxide generation due to higher antioxidative enzymes and reduced glutathione formation (GSH). Considering the obtained results, we can supplement the PGPR in combination to plants subjected to salt stress, prevent photosynthetic and growth reduction, and increase the yield of plants.

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 H 2 O 2 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., (© 2022 Scandinavian Plant Physiology Society.)

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.

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., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Sehar, Iqbal, Fatma, Rather, Albaqami and Khan.)

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 (H 2 S) 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 H 2 S is connected to the S-assimilation pathway, it is logical to consider the existence of a functional interplay between ethylene, H 2 S, and S in relation to heat stress tolerance. The present review focuses on the crosstalk between ethylene, H 2 S, and S to highlight their joint involvement in heat stress tolerance.

Nitric oxide (NO) and abscisic acid (ABA) play a significant role to combat abiotic stress. Application of 100 µM sodium nitroprusside (SNP, NO donor) or ABA alleviated heat stress effects on photosynthesis and growth of wheat ( Triticum aestivum L.) plants exposed to 40 °C for 6 h every day for 15 days. We have shown that ABA and NO synergistically interact to reduce the heat stress effects on photosynthesis and growth via reducing the content of H 2 O 2 and thiobarbituric acid reactive substances (TBARS), as well as maximizing osmolytes production and the activity and expression of antioxidant enzymes. The inhibition of NO and ABA using c-PTIO (2-4 carboxyphenyl-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide) and fluridone (Flu), respectively, reduced the osmolyte and antioxidant metabolism and heat stress tolerance. The inhibition of NO significantly reduced the ABA-induced osmolytes and antioxidant metabolism, exhibiting that the function of ABA in the alleviation of heat stress was NO dependent and can be enhanced with NO supplementation.Thus, regulating the activity and expression of antioxidant enzymes together with osmolytes production could act as a possible strategy for heat tolerance.

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 H 2 O 2 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 H 2 O 2 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.

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., (Copyright © 2021 The Authors. Published by Elsevier Inc. All rights reserved.)

Photosynthesis is a pivotal process that determines the synthesis of carbohydrates required for sustaining growth under normal or stress situation. Stress exposure reduces the photosynthetic potential owing to the excess synthesis of reactive oxygen species that disturb the proper functioning of photosynthetic apparatus. This decreased photosynthesis is associated with disturbances in carbohydrate metabolism resulting in reduced growth under stress. We evaluated the importance of melatonin in reducing heat stress-induced severity in wheat ( Triticum aestivum L.) plants. The plants were subjected to 25 °C (optimum temperature) or 40 °C (heat stress) for 15 days at 6 h time duration and then developed the plants for 30 days. Heat stress led to oxidative stress with increased production of thiobarbituric acid reactive substances (TBARS) and hydrogen peroxide (H 2 O 2 ) content and reduced accrual of total soluble sugars, starch and carbohydrate metabolism enzymes which were reflected in reduced photosynthesis. Application of melatonin not only reduced oxidative stress through lowering TBARS and H 2 O 2 content, augmenting the activity of antioxidative enzymes but also increased the photosynthesis in plant and carbohydrate metabolism that was needed to provide energy and carbon skeleton to the developing plant under stress. However, the increase in these parameters with melatonin was mediated via hydrogen sulfide (H 2 S), as the inhibition of H 2 S by hypotaurine (HT; H 2 S scavenger) reversed the ameliorative effect of melatonin. This suggests a crosstalk of melatonin and H 2 S in protecting heat stress-induced photosynthetic inhibition via regulation of carbohydrate metabolism.

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 (H 2 O 2 ), hydrogen sulfide (H 2 S), antioxidants, proline, glycine betaine, calcium (Ca 2+ ) 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., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The application of 10 µM methyl jasmonate (MeJA) for the protection of wheat ( Triticum aestivum L.) photosystem II (PS II) against heat stress (HS) was studied. Heat stress was induced at 42 °C to established plants, which were then recovered at 25 °C and monitored during their growth for the study duration. Application of MeJA resulted in increased enzymatic antioxidant activity that reduced the content of hydrogen peroxide (H 2 O 2 ) and thiobarbituric acid reactive substances (TBARS) and enhanced the photosynthetic efficiency. Exogenous MeJA had a beneficial effect on chlorophyll fluorescence under HS and enhanced the pigment system (PS) II system, as observed in a JIP-test, a new tool for chlorophyll fluorescence induction curve. Exogenous MeJA improved the quantum yield of electron transport (ET o /CS) as well as electron transport flux for each reaction center (ET 0 /RC). However, the specific energy fluxes per reaction center (RC), i.e., TR 0 /RC (trapping) and DI 0 /RC (dissipation), were reduced by MeJA. These results indicate that MeJA affects the efficiency of PS II by stabilizing the D1 protein, increasing its abundance, and enhancing the expression of the psb A and psb B genes under HS, which encode proteins of the PS II core RC complex. Thus, MeJA is a potential tool to protect PS II and D1 protein in wheat plants under HS and to accelerate the recovery of the photosynthetic capacity.

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 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.

Drought has been highly prevalent around the world especially in Sub-Saharan Africa and South-East Asian countries. Consistent climatic instabilities and unpredictable rainfall patterns are further worsening the situation. Rice is a C 3 staple cereal and an important food crop for the majority of the world's population and drought stress is one of the major growth retarding threats for rice that slashes down grain quality and yield. Drought deteriorates rice productivity and induces various acclimation responses that aids in stress mitigation. However, the complexity of traits associated with drought tolerance has made the understanding of drought stress-induced responses in rice a challenging process. An integrative understanding based on physiological adaptations, omics, transgenic and molecular breeding approaches successively backed up to developing drought stress-tolerant rice. The review represents a step forward to develop drought-resilient rice plants by exploiting the knowledge that collaborates with omics-based developments with integrative efforts to ensure the compilation of all the possible strategies undertaken to develop drought stress-tolerant rice., (© 2020 Scandinavian Plant Physiology Society.)

This study explored the interactive effect of ethephon (2-chloroethyl phosphonic acid; an ethylene source) and sulfur (S) in regulating the antioxidant system and ABA content and in maintaining stomatal responses, chloroplast structure, and photosynthetic performance of mustard plants ( Brassica juncea L. Czern.) grown under 100 mM NaCl stress. The treatment of ethephon (200 µL L -1 ) and S (200 mg S kg -1 soil) together markedly improved the activity of enzymatic and non-enzymatic components of the ascorbate-glutathione (AsA-GSH) cycle, resulting in declined oxidative stress through lesser content of sodium (Na + ) ion and hydrogen peroxide (H 2 O 2 ) in salt-stressed plants . These changes promoted the development of chloroplast thylakoids and photosynthetic performance under salt stress. Ethephon + S also reduced abscisic acid (ABA) accumulation in guard cell, leading to maximal stomatal conductance under salt stress. The inhibition of ethylene action by norbornadiene (NBD) in salt- plus non-stressed treated plants increased ABA and H 2 O 2 contents, and reduced stomatal opening, suggesting the involvement of ethephon and S in regulating stomatal conductance. These findings suggest that ethephon and S modulate antioxidant system and ABA accumulation in guard cells, controlling stomatal conductance, and the structure and efficiency of the photosynthetic apparatus in plants under salt stress.

The involvement of nitric oxide (NO) and hydrogen sulfide (H 2 S) in countermanding heat-inhibited photosynthetic features were studied in wheat ( Triticum aestivum L.). Heat stress (HS) was employed at 40 °C after establishment for 6 h daily, and then plants were allowed to recover at 25 °C and grown for 30 days. Glucose (Glc) content increased under HS and repressed plant photosynthetic ability, but the application of sodium nitroprusside (SNP, as NO donor) either alone or with sodium hydrosulfide (NaHS, as H 2 S donor) reduced Glc-mediated photosynthetic suppression by enhancing ascorbate-glutathione (AsA-GSH) metabolism and antioxidant system, which reduced oxidative stress with decreased H 2 O 2 and TBARS content. Oxidative stress reduction or inhibiting Glc repression was maximum with combined SNP and NaHS treatment, which was substantiated by 2-4-carboxyphenyl-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide (cPTIO) and hypotaurine (HT), scavengers for NO and H 2 S, respectively. The scavenge of H 2 S reduced NO-mediated alleviation of HS suggesting of its downstream action in NO-mediated heat-tolerance. However, a simultaneous decrease of both (NO and H 2 S) led to higher Glc-mediated repression of photosynthesis and oxidative stress in terms of increased H 2 O 2 content that was comparable to HS plants. Thus, NO and H 2 S cooperate to enhance photosynthesis under HS by reducing H 2 O 2 -induced oxidative stress and excess Glc-mediated photosynthetic suppression.

Gamma-aminobutyric acid (GABA) is a newly recognized signaling molecule participating in physiological processes, growth, and development of plants under optimal and stressful environments. In the present reported research, we investigated the role of GABA in imparting salt stress tolerance in wheat (Triticum aestivum L.). Exposure of wheat plants to 100 mM NaCl resulted in increased oxidative stress, glucose content, nitric oxide (NO) production together with reduced growth and photosynthetic traits of plants. Contrarily, GABA application improved nitrogen (N) metabolism, sulfur (S) assimilation, ion homeostasis, growth and photosynthesis under salt stress. Additionally, GABA mitigated oxidative stress induced by salt stress with the increased ascorbate-glutathione cycle and proline metabolism. The study with NO inhibitor, c-PTIO [2-(4-carboxyphenyl)-4,4,5,5-tetramethylimidazoline-1-oxy-3-oxide] in GABA experiment suggested that the impact of GABA on improvement of growth and photosynthesis under salt stress was mediated by NO and influenced N and S assimilation and antioxidant systems. The results suggested that the GABA has a significant potential in reversing the salt stress response in wheat plants, and GABA-mediated signals are manifested through NO., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Salinity is a serious threat to plant growth and development worldwide reducing agricultural productivity each year. Ethylene is an important phytohormone that affects plants performance under normal and abiotic stress conditions. In this study, role of ethylene was investigated in mitigating salinity stress (100 mM NaCl) effects on photosynthesis in mustard plants subjected to different nitrogen (N; 5 and 10 mM) levels. Plants under salinity stress exhibited marked increase in proline and reduced glutathione (GSH) content and activity of antioxidant enzymes. Nitrogen supplementation at 10 mM was better than 200 µl l -1 ethephon treatment under no stress. However, under salinity stress, both N and ethephon were equally effective. The combined application of 10 mM N and ethephon to salinity stressed plants produced greatest increase in photosynthesis by increasing proline and antioxidant metabolism. Ethylene evolution was high under salinity stress, but treatment of 10 mM N and 200 µl l -1 ethephon greatly decreased ethylene evolution that was equivalent to the 10 mM N treatment alone. This concentration of ethylene decreased the oxidative stress and increased the photosynthetic nitrogen use efficiency (NUE) maximally to increase photosynthesis. The use of ethylene action inhibitor, norbornadiene (NBD) showed reduction in ethylene mediated effects in alleviating salinity. Norbornadiene decreased the photosynthetic-NUE, proline and GSH content that resulted in decrease in photosynthesis under salinity stress. This study indicated that ethylene regulated the proline and antioxidant metabolism under salinity stress to increase photosynthetic functions of mustard grown with low and optimum N. The modulation of ethylene could be adopted in agricultural practices to increase photosynthesis under salinity stress.

The complex juvenile/maturity transition during a plant's life cycle includes growth, reproduction, and senescence of its fundamental organs: leaves, flowers, and fruits. Growth and senescence of leaves, flowers, and fruits involve several genetic networks where the phytohormone ethylene plays a key role, together with other hormones, integrating different signals and allowing the onset of conditions favorable for stage progression, reproductive success and organ longevity. Changes in ethylene level, its perception, and the hormonal crosstalk directly or indirectly regulate the lifespan of plants. The present review focused on ethylene's role in the development and senescence processes in leaves, flowers and fruits, paying special attention to the complex networks of ethylene crosstalk with other hormones. Moreover, aspects with limited information have been highlighted for future research, extending our understanding on the importance of ethylene during growth and senescence and boosting future research with the aim to improve the qualitative and quantitative traits of crops.

Ethylene is a plant hormone involved in several physiological processes and regulates the plant development during the whole life. Stressful conditions usually activate ethylene biosynthesis and signaling in plants. The availability of nutrients, shortage or excess, influences plant metabolism and ethylene plays an important role in plant adaptation under suboptimal conditions. Among the plant nutrients, the nitrogen (N) is one the most important mineral element required for plant growth and development. The availability of N significantly influences plant metabolism, including ethylene biology. The interaction between ethylene and N affects several physiological processes such as leaf gas exchanges, roots architecture, leaf, fruits, and flowers development. Low plant N use efficiency (NUE) leads to N loss and N deprivation, which affect ethylene biosynthesis and tissues sensitivity, inducing cell damage and ultimately lysis. Plants may respond differently to N availability balancing ethylene production through its signaling network. This review discusses the recent advances in the interaction between N availability and ethylene at whole plant and different organ levels, and explores how N availability induces ethylene biology and plant responses. Exogenously applied ethylene seems to cope the stress conditions and improves plant physiological performance. This can be explained considering the expression of ethylene biosynthesis and signaling genes under different N availability. A greater understanding of the regulation of N by means of ethylene modulation may help to increase NUE and directly influence crop productivity under conditions of limited N availability, leading to positive effects on the environment. Moreover, efforts should be focused on the effect of N deficiency or excess in fruit trees, where ethylene can have detrimental effects especially during postharvest.

Proline content and ethylene production have been shown to be involved in salt tolerance mechanisms in plants. To assess the role of nitrogen (N) in the protection of photosynthesis under salt stress, the effect of N (0, 5, 10, 20 mM) on proline and ethylene was studied in mustard (Brassica juncea). Sufficient N (10 mM) optimized proline production under non-saline conditions through an increase in proline-metabolizing enzymes, leading to osmotic balance and protection of photosynthesis through optimal ethylene production. Excess N (20 mM), in the absence of salt stress, inhibited photosynthesis and caused higher ethylene evolution but lower proline production compared to sufficient N. In contrast, under salt stress with an increased demand for N, excess N optimized ethylene production, which regulates the proline content resulting in recovered photosynthesis. The effect of excess N on photosynthesis under salt stress was further substantiated by the application of the ethylene biosynthesis inhibitor, 1-aminoethoxy vinylglycine (AVG), which inhibited proline production and photosynthesis. Without salt stress, AVG promoted photosynthesis in plants receiving excess N by inhibiting stress ethylene production. The results suggest that a regulatory interaction exists between ethylene, proline and N for salt tolerance. Nitrogen differentially regulates proline production and ethylene formation to alleviate the adverse effect of salinity on photosynthesis in mustard., (Copyright © 2015 Elsevier GmbH. All rights reserved.)

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., (© 2014 Scandinavian Plant Physiology Society.)

The plant hormone ethylene is involved in many physiological processes, including plant growth, development and senescence. Ethylene also plays a pivotal role in plant response or adaptation under biotic and abiotic stress conditions. In plants, ethylene production often enhances the tolerance to sub-optimal environmental conditions. This role is particularly important from both ecological and agricultural point of views. Among the abiotic stresses, the role of ethylene in plants under nutrient stress conditions has not been completely investigated. In literature few reports are available on the interaction among ethylene and macro- or micro-nutrients. However, the published works clearly demonstrated that several mineral nutrients largely affect ethylene biosynthesis and perception with a strong influence on plant physiology. The aim of this review is to revisit the old findings and recent advances of knowledge regarding the sub-optimal nutrient conditions on the effect of ethylene biosynthesis and perception in plants. The effect of deficiency or excess of the single macronutrient or micronutrient on the ethylene pathway and plant responses are reviewed and discussed. The synergistic and antagonist effect of the different mineral nutrients on ethylene plant responses is critically analyzed. Moreover, this review highlights the status of information between nutritional stresses and plant response, emphasizing the topics that should be further investigated., (Copyright © 2013 Elsevier Masson SAS. All rights reserved.)

We investigated the potential of salicylic acid (SA) in alleviating the adverse effects of heat stress on photosynthesis in wheat (Triticum aestivum L.) cv WH 711. Activity of ribulose 1,5-bisphosphate carboxylase (Rubisco), photosynthetic-nitrogen use efficiency (NUE), and net photosynthesis decreased in plants subjected to heat stress (40 °C for 6 h), but proline metabolism increased. SA treatment (0.5 mM) alleviated heat stress by increasing proline production through the increase in γ-glutamyl kinase (GK) and decrease in proline oxidase (PROX) activity, resulting in promotion of osmotic potential and water potential necessary for maintaining photosynthetic activity. Together with this, SA treatment restricted the ethylene formation in heat-stressed plants to optimal range by inhibiting activity of 1-aminocyclopropane carboxylic acid (ACC) synthase (ACS). This resulted in improved proline metabolism, N assimilation and photosynthesis. The results suggest that SA interacts with proline metabolism and ethylene formation to alleviate the adverse effects of heat stress on photosynthesis in wheat.

Sulfur (S) deficiency is prevailing all over the world and becoming an important issue for crop improvement through maximising its utilization efficiency by plants for sustainable agriculture. Its interaction with other regulatory molecules in plants is necessary to improve our understanding on its role under changing environment. Our knowledge on the influence of S on ethylene signaling is meagre although it is a constituent of cysteine (Cys) required for the synthesis of reduced glutathione (GSH) and S-adenosyl methionine (SAM), a precursor of ethylene biosynthesis. Thus, there may be an interaction between S assimilation, ethylene signaling and plant responses under optimal and stressful environmental conditions. The present review emphasizes that responses of plants to S involve ethylene action. This evaluation will provide an insight into the details of interactive role of S and ethylene signaling in regulating plant processes and prove profitable for developing sustainability under changing environmental conditions.

Ethylene controls photosynthesis and induces tolerance of plants to metal stress. However, little is known about the interaction between ethylene, photosynthesis and sulfur (S) availability under cadmium (Cd) stress. Recently, we reported that ethylene controls photosynthesis by increasing glutathione (GSH) synthesis with sufficient-S availability under Cd stress. Plants treated with Cd were less sensitive to ethylene and showed photosynthetic inhibition. Ethylene sensitivity of plants was increased with exogenously-sourced ethylene or with sufficient-S application resulting in induced GSH synthesis and alleviation of photosynthetic inhibition by Cd. In this addendum we present some additional data indicating that ethylene regulates photosynthesis by reducing glucose (Glc) sensitivity, thus reducing the Glc-mediated photosynthetic repression.

Sulphur (S) assimilation leads to the formation of glutathione (GSH) and alleviation of cadmium (Cd) stress. GSH is synthesized from its immediate metabolite cysteine, which also serves as a metabolite for ethylene formation through S-adenosyl methionine. To assess the role of ethylene in S-induced alleviation of Cd stress on photosynthesis, the effects of S or ethephon (ethylene source) on GSH and ethylene were examined in mustard (Brassica juncea L. cv. Varuna). Sufficient-S at 100 mg S kg(-1) soil alleviated Cd-induced photosynthetic inhibition more than excess-S (200 mg S kg(-1) soil) via ethylene by increased GSH. Under Cd stress, plants were less sensitive to ethylene, despite high ethylene evolution, and showed photosynthetic inhibition. Ethylene sensitivity of plants increased with ethephon or sufficient-S, triggering the induction of an antioxidant system, and leading to increased photosynthesis even under Cd stress. The effects of ethephon and S under Cd stress were similar. The effects of S were reversed by ethylene biosynthesis inhibitor, aminoethoxyvinylglycine (AVG), suggesting that ethylene plays an important role in S-induced alleviation of Cd stress on photosynthesis., (© 2011 Blackwell Publishing Ltd.)

In order to ascertain the stomatal and photosynthetic responses of mustard to ethylene under varying N availability, photosynthetic characteristics of mustard grown with optimal (80 mg N kg(-1) soil) or low (40 mg N kg(-1) soil) N were studied after the application of an ethylene-releasing compound, ethephon (2-chloroethyl phosphonic acid) at 40 days after sowing (DAS). The availability of N influenced ethylene evolution and affected stomatal conductance and photosynthesis. The effect of ethylene was smaller under deficient N where plants contained higher glucose (Glc) sensitivity, despite high ethylene evolution even in the absence of ethephon, potentially because the plants were less sensitive to ethylene per se. Ethephon application at each level of N increased ethylene and decreased Glc sensitivity, which increased photosynthesis via its effect on the photosynthetic machinery and effects on stomatal conductance. Plants grown with sufficient-N and treated with 200 μl l(-1) ethephon exhibited optimal ethylene, the greatest stomatal conductance and photosynthesis, and growth. These plants made maximum use of available N and exhibited the highest nitrogen-use efficiency (NUE).

Salicylic acid (SA) is known to affect photosynthesis under normal conditions and induces tolerance in plants to biotic and abiotic stresses through influencing physiological processes. In this study, physiological processes were compared in salt-tolerant (Pusa Vishal) and salt-sensitive (T44) cultivars of mungbean and examined how much these processes were induced by SA treatment to alleviate decrease in photosynthesis under salt stress. Cultivar T44 accumulated higher leaf Na(+) and Cl(-) content and exhibited greater oxidative stress than Pusa Vishal. Activity of antioxidant enzymes, ascorbate peroxidase (APX) and glutathione reductase (GR) was greater in Pusa Vishal than T44. Contrarily, activity of superoxide dismutase (SOD) was greater in T44. The greater accumulation of leaf nitrogen and sulfur through higher activity of their assimilating enzymes, nitrate reductase (NR) and ATP-sulfurylase (ATPS) increased reduced glutathione (GSH) content more conspicuously in Pusa Vishal than T44. Application of 0.5 mM SA increased nitrogen and sulfur assimilation, GSH content and activity of APX and GR. This resulted in the increase in photosynthesis under non-saline condition and alleviated the decrease in photosynthesis under salt stress. It also helped in restricting Na(+) and Cl(-) content in leaf, and maintaining higher efficiency of PSII, photosynthetic N-use efficiency (NUE) and water relations in Pusa Vishal. However, application of 1.0 mM SA resulted in inhibitory effects. The effect of SA was more pronounced in Pusa Vishal than T44. These results indicate that SA application alleviates the salt-induced decrease in photosynthesis mainly through inducing the activity of NR and ATPS, and increasing antioxidant metabolism to a greater extent in Pusa Vishal than T44., (Copyright © 2010 Elsevier GmbH. All rights reserved.)