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.

Brassinosteroid modulates ethylene synthesis and antioxidant metabolism to protect rice (Oryza sativa) against heat stress-induced inhibition of source‒sink capacity and photosynthetic and growth attributes.

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.

Methyl jasmonate influences ethylene formation, defense systems, nutrient homeostasis and carbohydrate metabolism to alleviate arsenic-induced stress in rice (Oryza sativa).

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.

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.

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.

Palliating Salt Stress in Mustard through Plant-Growth-Promoting Rhizobacteria: Regulation of Secondary Metabolites, Osmolytes, Antioxidative Enzymes and Stress Ethylene.

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 H2O2 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.

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.

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.

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.

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.

Ethylene and Sulfur Coordinately Modulate the Antioxidant System and ABA Accumulation in Mustard Plants under Salt Stress.

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 (H2O2) 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 H2O2 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.

Ethephon increases photosynthetic-nitrogen use efficiency, proline and antioxidant metabolism to alleviate decrease in photosynthesis under salinity stress in mustard.

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.

Ethylene Role in Plant Growth, Development and Senescence: Interaction with Other Phytohormones.

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.

Role of ethylene in responses of plants to nitrogen availability.

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.

Nitrogen availability regulates proline and ethylene production and alleviates salinity stress in mustard (Brassica juncea).

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.