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

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

Phytoremediation of Potentially Toxic Elements: Role, Status and Concerns.

Environmental contamination with a myriad of potentially toxic elements (PTEs) is triggered by various natural and anthropogenic activities. However, the industrial revolution has increased the intensity of these hazardous elements and their concentration in the environment, which, in turn, could provoke potential ecological risks. Additionally, most PTEs pose a considerable nuisance to human beings and affect soil, aquatic organisms, and even nematodes and microbes. This comprehensive review aims to: (i) introduce potentially toxic elements; (ii) overview the major sources of PTEs in the major environmental compartments; (iii) briefly highlight the major impacts of PTEs on humans, plants, aquatic life, and the health of soil; (iv) appraise the major methods for tackling PTE-caused pollution; (v) discuss the concept and applications of the major eco-technological/green approaches (comprising phytoextraction, rhizofiltration, phytostabilization, phytovolatilization, and phytorestoration); (vi) highlight the role of microbes in phytoremediation under PTE stress; and (vii) enlighten the major role of genetic engineering in advancing the phytoremediation of varied PTEs. Overall, appropriate strategies must be developed in order to stop gene flow into wild species, and biosafety issues must be properly addressed. Additionally, consistent efforts should be undertaken to tackle the major issues (e.g., risk estimation, understanding, acceptance and feasibility) in order to guarantee the successful implementation of phytoremediation programs, raise awareness of this green technology among laymen, and to strengthen networking among scientists, stakeholders, industrialists, governments and non-government organizations.

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.

Exogenously Applied Rohitukine Inhibits Photosynthetic Processes, Growth and Induces Antioxidant Defense System in Arabidopsis thaliana.

The secondary metabolite rohitukine has been reported in only a few plant species, including Schumanniophyton magnificum, S. problematicum, Amoora rohituka, Dysoxylum acutangulum and D. gotadhora. It has several biological activities, such as anticancer, anti-inflammatory, antiadipogenic, immunomodulatory, gastroprotective, anti-implantation, antidyslipidemic, anti-arthritic and anti-fertility properties. However, the ecological and physiological roles of rohitukine in parent plants have yet to be explored. Here for the first time, we tried to decipher the physiological effect of rohitukine isolated from D. gotadhora on the model system Arabidopsis thaliana. Application of 0.25 mM and 0.5 mM rohitukine concentrations moderately affected the growth of A. thaliana, whereas a remarkable decrease in growth and the alteration of various morphological, physiological and biochemical mechanisms were observed in plants that received 1.0 mM of rohitukine as compared to the untreated control. A. thaliana showed considerable dose-dependent decreases in leaf area, fresh weight and dry weight when sprayed with 0.25 mM, 0.5 mM and 1.0 mM of rohitukine. Rohitukine exposure resulted in the disruption of photosynthesis, photosystem II (PSII) activity and degradation of chlorophyll content in A. thaliana. It also triggered oxidative stress in visualized tissues through antioxidant enzyme activity and the expression levels of key genes involved in the antioxidant system, such as superoxide dismutase (SOD), peroxidase (POD) and ascorbate peroxidase (APX). Rohitukine-induced changes in levels of metabolites (amino acids, sugars, organic acids, etc.) were also assessed. In light of these results, we discuss (i) the likely ecological importance of rohitukine in parent plants as well as (ii) the comparison of responses to rohitukine treatment in plants and mammals.

Exogenously-applied L-glutamic acid protects photosynthetic functions and enhances arsenic tolerance through increased nitrogen assimilation and antioxidant capacity in rice (Oryza sativa L.).

L-Glutamic acid (Glu) is used as an effective bio-stimulant to reduce arsenic (As) stress in plants. The role of Glu was studied in the protection of photosynthesis and growth of rice (Oryza sativa L. Japonica Type Taipie-309) plants grown with 50 µM As stress by studying the oxidative stress, photosynthetic and growth characteristics. Among the Glu concentrations (0, 2.5, 5 and 10 µM), 10 µM Glu maximally enhanced photosynthesis and growth parameters with the least cellular oxidative stress level. The supplementation of 10 µM Glu resulted in the reduced effects of As stress on gas exchange parameters, PSII activity and growth attributes through enhancement of antioxidant and proline metabolism. The enzymes of nitrogen (N) assimilation, such as nitrate reductase, nitrite reductase, glutamine synthetase and glutamate synthase were increased with Glu treatment under As stress. The Glu-induced metabolite synthesis showed the role of various metabolites in As stress responses. The role of Glu as a signalling molecule in reducing the adverse effects of As through accelerating the antioxidant enzymes, PSII activity, proline metabolism and nitrogen assimilation has been discussed.

Role of Jasmonates, Calcium, and Glutathione in Plants to Combat Abiotic Stresses Through Precise Signaling Cascade.

Plant growth regulators have an important role in various developmental processes during the life cycle of plants. They are involved in abiotic stress responses and tolerance. They have very well-developed capabilities to sense the changes in their external milieu and initiate an appropriate signaling cascade that leads to the activation of plant defense mechanisms. The plant defense system activation causes build-up of plant defense hormones like jasmonic acid (JA) and antioxidant systems like glutathione (GSH). Moreover, calcium (Ca2+) transients are also seen during abiotic stress conditions depicting the role of Ca2+ in alleviating abiotic stress as well. Therefore, these growth regulators tend to control plant growth under varying abiotic stresses by regulating its oxidative defense and detoxification system. This review highlights the role of Jasmonates, Calcium, and glutathione in abiotic stress tolerance and activation of possible novel interlinked signaling cascade between them. Further, phyto-hormone crosstalk with jasmonates, calcium and glutathione under abiotic stress conditions followed by brief insights on omics approaches is also elucidated.

Plant aquaporins: A frontward to make crop plants drought resistant.

Drought stress alters gene expression and causes cellular damage in crop plants. Drought inhibits photosynthesis by reducing the content and the activity of the photosynthetic carbon reduction cycle, ultimately decreasing the crop yield. The role of aquaporins (AQP) in improving the growth and adaptation of crop plants under drought stress is of importance. AQP form channels and control water transport in and out of the cells and are associated with drought tolerance mechanisms. The current review addresses: (1) the evolution of AQPs in plants, (2) the classification of plant AQPs, (3) the role of AQPs in drought alleviation in crop plants, and (4) the phytohormone crosstalk with AQPs in crops exposed to drought stress.

Hydrogen peroxide modulates activity and expression of antioxidant enzymes and protects photosynthetic activity from arsenic damage in rice (Oryza sativa L.).

We studied the role of H2O2 in the protection of photosynthesis from arsenic (As) damage in rice (Oryza sativa L.) by examining the antioxidant system, photosynthesis, and growth attributes. Among the As concentrations (0, 20, 30, 40 and 50 µM) tested, maximum oxidative stress and inhibition in photosynthesis and growth were found with 50 µM As. The application of 50 µM H2O2 resulted in alleviation of the adverse effects of 50 µM As on Pigment System (PS) II activity, photosynthesis, and growth. Hydrogen peroxide supplementation induced the activity of superoxide dismutase (SOD), ascorbate peroxidase (APX) and glutathione reductase (GR) and increased reduced glutathione (GSH) content and proline metabolism. The expression of SOD and APX, PSBA and PSBB was induced in the presence of H2O2 to alleviate the As damage to PS II and maintain photosynthetic activity. The role of H2O2 as a signaling molecule is shown in the protection of photosynthetic activity in rice from As toxicity through regulation on the activity and the expression of antioxidant enzymes.

Physiological, Biochemical and Reproductive Studies on Valeriana wallichii, a Critically Endangered Medicinal Plant of the Himalayan Region Grown under In-Situ and Ex-Situ Conditions.

Valeriana wallichii, a perennial herb belonging to family Valerianaceae, is an important medicinal herb of the Himalayan region. The incessant exploitation of nature for meeting the demands of the pharmaceutical industry has put unbearable pressure on its natural habitats. A study on its physiological, biochemical, growth and reproductive attributes was planned. Physiological study revealed that ex-situ (outside their natural habitat) populations faced severe stress as compared to in-situ (natural habitat) plants. The difference in the performance of these habitat plants was related to superoxide and H2O2 in the leaves. Photosynthetic attributes were increased in in-situ populations. Proline content and its biosynthetic enzymes ornithine aminotransferase, and pyrroline-5-carboxylate reductase showed an increase in ex-situ plants; proline oxidase decreased. Glucose-6-phosphate dehydrogenase, shikimic acid dehydrogenese, phenylalanine lyase, and flavonoids content showed an increment in ex-situ plants. Antioxidants enzyme superoxide dismutase, catalase, ascorbate peroxidase and reduced glutathione showed an increment in ex-situ conditions. Growth and reproductive attributes were more in ex-situ plants. The observations made are suggestive that a comprehensive conservation programme involving in-situ as well as ex-situ strategies will be effective for the conservation and long term survival of the species.

Nitric oxide-mediated regulation of oxidative stress in plants under metal stress: a review on molecular and biochemical aspects.

Given their sessile nature, plants continuously face unfavorable conditions throughout their life cycle, including water scarcity, extreme temperatures and soil pollution. Among all, metal(loid)s are one of the main classes of contaminants worldwide, posing a serious threat to plant growth and development. When in excess, metals which include both essential and non-essential elements, quickly become phytotoxic, inducing the occurrence of oxidative stress. In this way, in order to ensure food production and safety, attempts to enhance plant tolerance to metal(loid)s are urgently needed. Nitric oxide (NO) is recognized as a signaling molecule, highly involved in multiple physiological events, like the response of plants to abiotic stress. Thus, substantial efforts have been made to assess NO potential in alleviating metal-induced oxidative stress in plants. In this review, an updated overview of NO-mediated protection against metal toxicity is provided. After carefully reviewing NO biosynthetic pathways, focus was given to the interaction between NO and the redox homeostasis followed by photosynthetic performance of plants under metal excess.

Nitric oxide signaling and its crosstalk with other plant growth regulators in plant responses to abiotic stress.

Nitric oxide (NO) is a free radical molecule involved in an array of functions under physiological and adverse environmental conditions. As other free radical molecules, NO biological action depends on its cellular concentration, acting as a signal molecule when produced at low concentration or resulting in cellular damage when produced at sufficiently high levels to trigger nitro-oxidative stress. Over the last decade, significant progress has been made in characterizing NO metabolism and action mechanism, revealing that diverse biosynthetic routes can generate this free radical in plants and its action mainly occurs through posttranslational modification (nitration and S-nitrosylation) of target proteins. Intricate crosstalk networks between NO and other signaling molecules have been described involving phytohormones, other second messengers, and key transcription factors. This review will focus on our current understanding of NO interplay with phytohormones and other plant growth regulators under abiotic stress conditions.

Ethylene Potentiates Sulfur-Mediated Reversal of Cadmium Inhibited Photosynthetic Responses in Mustard.

The potential of exogenous ethylene and sulfur (S) in reversal of cadmium (Cd)-inhibited photosynthetic and growth responses in mustard (Brassica juncea L. cv. Pusa Jai Kisan) were studied. Plants grown with 50 µM Cd showed increased superoxide and H2O2 accumulation and lipid peroxidation together with increased activity of 1-aminocyclopropane carboxylic acid synthase (ACS) and ethylene production and inhibition of photosynthesis and growth. Application of 1 mM SO42- or 200 µL L-1 ethephon (ethylene source) influenced photosynthetic and growth performance equally in presence or absence of Cd. However, their combined application synergistically improved photosynthetic performance more in presence of Cd and reduced oxidative stress (lower superoxide and H2O2 accumulation) by decreasing ethylene and glucose sensitivity with the increase in cysteine and methionineand a non-proteinogenic thiol (reduced glutathione; GSH) contents. The central role of ethylene in potentiating S-mediated reversal of Cd-induced oxidative stress was evident with the use of ethylene action inhibitor, norbornadiene (NBD). The application of NBD resulted in decreased thiol production and photosynthetic responses. This suggests that ethylene promotes the effects of S in reversal of adverse effects of Cd, and thus, ethylene modulation may be considered as potential tool to substantiate the S effects in reversal of Cd inhibited photosynthesis and growth in mustard.

Involvement of ethylene in gibberellic acid-induced sulfur assimilation, photosynthetic responses, and alleviation of cadmium stress in mustard.

The role of gibberellic acid (GA) or sulfur (S) in stimulation of photosynthesis is known. However, information on the involvement of ethylene in GA-induced photosynthetic responses and cadmium (Cd) tolerance is lacking. This work shows that ethylene is involved in S-assimilation, photosynthetic responses and alleviation of Cd stress by GA in mustard (Brassica juncea L.). Plants grown with 200 mg Cd kg(-1) soil were less responsive to ethylene despite high ethylene evolution and showed photosynthetic inhibition. Plants receiving 10 µM GA spraying plus 100 mg S kg(-1) soil supplementation exhibited increased S-assimilation and photosynthetic responses under Cd stress. Application of GA plus S decreased oxidative stress of plants grown with Cd and limited stress ethylene formation to the range suitable for promoting sulfur use efficiency (SUE), glutathione (GSH) production and photosynthesis. The role of ethylene in GA-induced S-assimilation and reversal of photosynthetic inhibition by Cd was substantiated by inhibiting ethylene biosynthesis with the use of aminoethoxyvinylglycine (AVG). The suppression of S-assimilation and photosynthetic responses by inhibiting ethylene in GA plus S treated plants under Cd stress indicated the involvement of ethylene in GA-induced S-assimilation and Cd stress alleviation. The outcome of the study is important to unravel the interaction between GA and ethylene and their role in Cd tolerance in plants.

Hydrogen Peroxide Alleviates Nickel-Inhibited Photosynthetic Responses through Increase in Use-Efficiency of Nitrogen and Sulfur, and Glutathione Production in Mustard.

The response of two mustard (Brassica juncea L.) cultivars differing in photosynthetic capacity to different concentrations of hydrogen peroxide (H2O2) or nickel (Ni) was evaluated. Further, the effect of H2O2 on photosynthetic responses of the mustard cultivars grown with or without Ni stress was studied. Application of 50 µM H2O2 increased photosynthesis and growth more prominently in high photosynthetic capacity cultivar (Varuna) than low photosynthetic capacity cultivar (RH30) grown without Ni stress. The H2O2 application also resulted in alleviation of photosynthetic inhibition induced by 200 mg Ni kg(-1) soil through increased photosynthetic nitrogen-use efficiency (NUE), sulfur-use efficiency (SUE), and glutathione (GSH) reduced production together with decreased lipid peroxidation and electrolyte leakage in both the cultivars. However, the effect of H2O2 was more pronounced in Varuna than RH30. The greater increase in photosynthetic-NUE and SUE and GSH production with H2O2 in Varuna resulted from higher increase in activity of nitrogen (N) and sulfur (S) assimilation enzymes, nitrate reductase and ATP-sulfurylase, respectively resulting in enhanced N and S assimilation. The increased N and S content contributed to the higher activity of ribulose-1,5-bisphosphate carboxylase under Ni stress. Application of H2O2 also regulated PS II activity and stomatal movement under Ni stress for maintaining higher photosynthetic potential in Varuna. Thus, H2O2 may be considered as a potential signaling molecule for augmenting photosynthetic potential of mustard plants under optimal and Ni stress conditions. It alleviates Ni stress through the regulation of stomatal and non-stomotal limitations, and photosynthetic-NUE and -SUE and GSH production.

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

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

Minimising toxicity of cadmium in plants--role of plant growth regulators.

A range of man-made activities promote the enrichment of world-wide agricultural soils with a myriad of chemical pollutants including cadmium (Cd). Owing to its significant toxic consequences in plants, Cd has been one of extensively studied metals. However, sustainable strategies for minimising Cd impacts in plants have been little explored. Plant growth regulators (PGRs) are known for their role in the regulation of numerous developmental processes. Among major PGRs, plant hormones (such as auxins, gibberellins, cytokinins, abscisic acid, jasmonic acid, ethylene and salicylic acid), nitric oxide (a gaseous signalling molecule), brassinosteroids (steroidal phytohormones) and polyamines (group of phytohormone-like aliphatic amine natural compounds with aliphatic nitrogen structure) have gained attention by agronomist and physiologist as a sustainable media to induce tolerance in abiotic-stressed plants. Considering recent literature, this paper: (a) overviews Cd status in soil and its toxicity in plants, (b) introduces major PGRs and overviews their signalling in Cd-exposed plants, (c) appraises mechanisms potentially involved in PGR-mediated enhanced plant tolerance to Cd and (d) highlights key aspects so far unexplored in the subject area.

Selenium and sulfur influence ethylene formation and alleviate cadmium-induced oxidative stress by improving proline and glutathione production in wheat.

We have studied the influence of selenium (Se) and sulfur (S) in the protection of photosynthetic capacity of wheat (Triticum aestivum) against cadmium (Cd) stress. The involvement of ethylene and its interaction with proline and antioxidant metabolism in the tolerance of plants to Cd stress was evaluated. Application of Se or S alleviated Cd-induced oxidative stress by increasing proline accumulation as a result of increased activity of glutamyl kinase (GK) and decreased activity of proline oxidase (PROX). These nutrients also induced the activity of ATP-sulfurylase and serine acetyl transferase and the content of cysteine (Cys), a precursor for the synthesis of both reduced glutathione (GSH) and ethylene. Further, application of Se and S to plants under Cd stress reduced ethylene level and increased the activity of glutathione reductase (GR) and glutathione peroxidase (GPX), reduced oxidative stress and improved photosynthesis and growth. The involvement of ethylene in Se and S-mediated alleviation of Cd stress was substantiated with the use of ethylene biosynthesis inhibitor aminoethoxyvinylglycine (AVG). The use of AVG reversed the effects of Se and S on ethylene, content of proline and GSH and photosynthesis. The results suggested that Se and S both reversed Cd-induced oxidative stress by regulating ethylene formation, proline and GSH metabolism. Thus, Se or S-induced regulatory interaction between ethylene and proline and GSH metabolism may be used for the reversal of Cd-induced oxidative stress.

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

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

Cross-talk between sulfur assimilation and ethylene signaling in plants.

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.