Involvement of salicylic acid and nitric oxide in protective reactions of wheat under the influence of heavy metals

This article studies the effect of salicylic acid (SA) and nitric oxide (NO) on Triticum aestivum L. wheat plants exposed to the influence of high concentrations of copper and zinc compounds. It is shown that heavy metals (HMs) caused a decrease in the growth parameters in the overground and underground plant parts and contributed to a sharp deterioration in the energy balance and the situation regarding oxidative stress. SA and NO exerted a protective effect, which was expressed in the increased ability to accumulate shoot and root mass, stabilize the energy balance, and reduced lipid peroxidation.     

Salicylic acid and nitric oxide alleviate osmotic stress in wheat (Triticum aestivum L.) seedlings

Salicylic acid (SA) and nitric oxide (NO) are reported to alleviate the damaging effects of stress in plants rather similarly when applied at appropriate low concentrations. An experiment was therefore conducted to study the impact of SA, sodium nitroprusside (SNP; as NO donor), and methylene blue (MB; as a guanylate cyclase inhibitor) on wheat seedling performance under osmotic stress. Osmotic stress significantly reduced shoot fresh weight (SFW), chlorophyll contents (Chla, Chlb, total Chl), and membrane stability index (MSI) and also increased malondialdehyde (MDA) level, lipoxygenase (LOX) activity, and hydrogen peroxide production. Moreover, enzymatic antioxidant activities including superoxide dismutase, guaiacol peroxidase, and glutathione reductase activity were enhanced under osmotic stress. On the contrary, SA or SNP pretreatment reduced the damaging effects of osmotic stress by further enhancing the antioxidant activities that led to increased SFW, Chl, and MSI and reduced MDA level and LOX activity. However, pretreatment of plants with MB reversed or reduced the protective effects of SA and SNP suggesting that the protective effects were likely attributed to NO signaling. Therefore, NO may act as downstream of SA signaling in reduction of induced oxidative damage in wheat seedlings.     

Effect of Increasing Concentrations of Heavy Metals on the Growth of Barley and Wheat Seedlings

The tolerance to increasing doses of lead and cadmium salts on the growth and survival of barley (Hordeum vulgare L.) and wheat (Triticum aestivum L.) seedlings were studied. Seedlings grown under controlled conditions were treated with dilute (0.001–0.005 mM) solutions of either lead nitrate or cadmium bromide for 1, 4, or 7 days. Subsequently, they were incubated for 7 days in solutions of the same compounds, but at sublethal or lethal concentrations (0.05–10 mM). Plant pretreatment with low concentrations of heavy metals induced an increase in their tolerance to the metals, because pretreated plants could tolerate heavy metals at high concentrations. It is concluded that plant tolerance to increasing concentrations of heavy metals is related to the activation of protective and adaptive processes in their tissues.     

Exogenous treatment with salicylic acid attenuates cadmium toxicity in pea seedlings

The present study investigated the possible mediatory role of salicylic acid (SA) in protecting plants from cadmium (Cd) toxicity. The exposure of pea plants to increasing Cd concentrations (0.5, 1.0, 2.0 and 5.0 μM) during early stages of their establishment, caused a gradual decrease in shoot and root fresh weight accumulation, the rate of CO₂ fixation and the activity of ribulose-1,5-bisphosphate carboxylase (RuBPC, E.C. 4.1.1.39), the effect being most expressed at higher Cd concentrations. In vivo the excess of Cd-induced alterations in the redox cycling of oxygen-evolving centers and the assimilatory capacity of the pea leaves as revealed by changes in thermoluminescence emission after flash illumination. The levels of some important parameters associated with oxidative stress, namely lipid peroxidation, electrolyte leakage and proline production were increased. Seed pretreatment with SA alleviated the negative effect of Cd on growth, photosynthesis, carboxylation reactions, thermoluminescence characteristics and chlorophyll content, and led to decrease in oxidative injuries caused by Cd. The data suggest that the beneficial effect of SA during an earlier growth period could be related to avoidance of cumulative damage upon exposure to cadmium thus reducing the negative consequences of oxidative stress caused by heavy metal toxicity. In addition, the observed high endogenous levels of SA after treatment with Cd suggests that SA may act directly as an antioxidant to scavenge the reactive oxygen species and/or indirectly modulate redox balance through activation of antioxidant responses.Taken together these evidences could explain at some extend the protective role of SA on photochemical activity of chloroplast membranes and photosynthetic carboxylation reactions in Cd-stressed pea plants.     

Combined Effect of Salicylic Acid and Nitrogen Oxide Donor on Stress-Protective System of Wheat Plants under Drought Conditions

Presowing treatment of wheat (Triticum aestivum L.) seeds with 10 or 100 μM salicylic acid (SA) reduced the inhibition of 14-day-old plant growth under soil drought. The same effect was caused by the spraying of 7-day-old seedlings with 0.5 or 2 mM nitrogen oxide donor (sodium nitroprusside, SNP) before drought. The protective effect was enhanced by the combination of seed treatment with 10 μM SA and plant spraying with 0.5 mM SNP, while their combinations in higher concentrations caused weaker effects. SA treatment in both concentrations and 0.5 mM SNP under drought conditions increased the antioxidant enzyme activity (superoxide dismutase, catalase, and guaiacol peroxidase) in leaves. This effect was especially significant when 10 μM SA was combined with 0.5 mM SNP. Spraying with 2 mM SNP and its combination with seed presowing with 100 μM SA did not significantly change the antioxidant enzyme activity; however, the proline content in the leaves increased. It is concluded that the SA stress-protective action on plants can be modified with exogenous nitrogen oxide.     

Cross-talk between salicylic acid and NaCl-generated reactive oxygen species and nitric oxide in tomato during acclimation to high salinity

Hydrogen peroxide (H₂O₂) and nitric oxide (NO) generated by salicylic acid (SA) are considered to be functional links of cross‐tolerance to various stressors. SA‐stimulated pre‐adaptation state was beneficial in the acclimation to subsequent salt stress in tomato (Solanum lycopersicum cv. Rio Fuego). At the whole‐plant level, SA‐induced massive H₂O₂ accumulation only at high concentrations (10^?3–10^?2M), which later caused the death of plants. The excess accumulation of H₂O₂ as compared with plants exposed to 100 mM NaCl was not associated with salt stress response after SA pre‐treatments. In the root tips, 10^?3–10^?2M SA triggered the production of reactive oxygen species (ROS) and NO with a concomitant decline in the cell viability. Sublethal concentrations of SA, however, decreased the effect of salt stress on ROS and NO production in the root apex. The attenuation of oxidative stress because of high salinity occurred not only in pre‐adapted plants but also at cell level. When protoplasts prepared from control leaves were exposed to SA in the presence of 100 mM NaCl, the production of NO and ROS was much lower and the viability of the cells was higher than in salt‐treated samples. This suggests that, the cross‐talk of signalling pathways induced by SA and high salinity may occur at the level of ROS and NO production. Abscisic acid (ABA), polyamines and 1‐aminocyclopropane‐1‐carboxylic acid, the compounds accumulating in pre‐treated plants, enhanced the diphenylene iodonium‐sensitive ROS and NO levels, but, in contrast to others, ABA and putrescine preserved the viability of protoplasts.     

Effect of heavy metals on isoperoxidases of Wheat

The influence of increasing concentrations of copper, zinc, lead, nickel, chromium and cadmium on 14-day-old seedlings of wheat (Triticum aestivum L. cv. Vergina) was studied. Plants were grown in 1/10 strength Rorison’s nutrient solution with increasing concentrations of each of the metals added separately. The toxicity of metals depressed shoot growth but the most evident symptoms were on roots. The concentration of each metal which caused inhibition of root growth was chosen to study the influence of metals on isoperoxidases of wheat shoots. The concentrations employed did not alter the number of peroxidase bands but almost in all cases enhanced the intensities of bands of pH 4.0-4.2 and 5.0-5.4, while they decreased the intensities of bands of pH 4.2-4.6 and 5.4-6.5. The similar effects of the different heavy metals employed may suggest similarity in metal action on wheat isoperoxidases. The increased intensities of peroxidase bands may be considered as an indication of enhanced senescence caused by the heavy metal treatments. Generally, our results suggest that the heavy metals employed have caused complex changes on the multiple forms of peroxidases.     

Plant virus infection development as affected by heavy metal stress

The work was focused on the investigation of possible dependencies between the development of viral infection in plants and the presence of high heavy metal concentrations in soil. Field experiments have been conducted in order to study the development of systemic tobacco mosaic virus (TMV) infection in Lycopersicon esculentum L. cv. Miliana plants under effect of separate salts of heavy metals Cu, Zn and Pb deposited in soil. As it is shown, simultaneous effect of viral infection and heavy metals in tenfold maximum permissible concentration leads to decrease of total chlorophyll content in experiment plants mainly due to the degradation of chlorophyll a. The reduction of chlorophyll concentration under the combined influence of both stress factors was more serious comparing to the separate effect of every single factor. Plants' treatment with toxic concentrations of lead and zinc leaded to slight delay in the development of systemic TMV infection together with more than twofold increase of virus content in plants that may be an evidence of synergism between these heavy metal's and virus' effects. Contrary, copper although decreased total chlorophyll content but showed protective properties and significantly reduced amount of virus in plants.     

Effect of Salicylic Acid on the Activity of Antioxidant Enzymes in Wheat under Conditions of Salination

The effect of pretreatment with 0.05 mM salicylic acid (SA) on the activity of superoxide dismutase (SOD) and peroxidase in the roots of four-day-old seedlings of wheat (Triticum aestivum L.) was studied under conditions of salination. The level of the stress-induced accumulation of active oxygen species and, therefore, activities of SOD and peroxidase in seedlings pretreated with SA were significantly lower than in untreated seedlings, which indicates that these enzymes contribute to the protective effect of SA on plants under conditions of salination.     

Nitric oxide stimulates seed germination and counteracts the inhibitory effect of heavy metals and salinity on root growth of Lupinus luteus

Nitric oxide (NO) is a bioactive molecule, which in plants was found to function as prooxidant as well as antioxidant. In the present study, we found that NO donor sodium nitroprusside (SNP) stimulates seed germination and root growth of lupin (Lupinus luteus L. cv. Ventus). Seed germination is promoted at concentrations between 0.1 and 800 μM SNP in a dose-dependent manner. The stimulation was most pronounced after 18 and 24 h and ceased after 48 h of imbibition. The promoting effect of NO on seed germination persisted even in the presence of heavy metals (Pb, Cd) and sodium chloride. Pretreatment of lupin seedlings for 24 h with 10 μM SNP resulted in efficient reduction of the detrimental effect of the abiotic stressors on root growth and morphology. The inhibitory effect of heavy metals on root growth was accompanied by increased activity of superoxide dismutase (SOD, EC 1.15.1.1.), which in roots preincubated with SNP was significantly higher. Some changes in the activity of other antioxidant enzymes, peroxidase (POX, EC 1.11.1.7) and catalase (CAT, EC 1.11.1.6) were also detected. Using the superoxide anion (O₂^•–)-specific indicator dihydroethidium (DHE), we found intense DHE-derived fluorescence in heavy metal-stressed roots, whereas in those pretreated with SNP the fluorescence was very low, comparable to the level in unstressed roots. On the basis of the above data, we conclude that the protective effect of NO in stressed lupin roots may be at least partly due to the stimulation of SOD activity and/or direct scavenging of the superoxide anion.     

Effects of exogenously applied salicylic acid and putrescine alone and in combination with rhizobacteria on the phytoremediation of heavy metals and chickpea growth in sandy soil

The present attempt was made to study the role of exogenously applied salicylic acid (SA) and putrescine (Put) on the phytoremediation of heavy metals and on the growth parameters of chickpea grown in sandy soil. The SA and Put were applied alone as well as in combination with plant growth promoting rhizobacteria (PGPR). The PGPRs, isolated from the rhizosphere of chickpea, were characterized on the basis of colony morphology and biochemical traits through gram staining, catalase and oxidase tests, and identified by 16S-rRNA gene sequencing as Bacillus subtilis, Bacillus thuringiensis and Bacillus megaterium. The chickpea seeds were soaked in 24 h old fresh cultures of isolates for 2–3 h prior to sowing. The growth regulators (PGRs), SA and Put (150 mg/L), were applied to the seedlings as foliar spray at three-leaf stage. The result revealed that plants treated with SA and Put alone or in combination with PGPRs, significantly enhanced the accumulation of heavy metals in plant shoot. PGPR induces Ni accumulation in sensitive variety and Pb in both the varieties, the PGR in combination augment the bioremediation effects of PGPR and both sensitive and tolerant variety showed significant accumulation of Ni, Cd, and Pb. SA was more effective in accumulating Ni and Cd whereas, significant accumulation of Pb was recorded in Put. PGPRs further augmented the PGRs induced accumulation of heavy metals and macronutrients in chickpea shoot and in rhizosphere. SA increased the proline content of tolerant variety while decreasing the lipid peroxidation and proline content of the sensitive variety but decreased the stimulating effect of PGPR in proline production. Interactive effects of PGPR and PGRs are recommended for inducing phytoremediation in chickpea plants under drought stress.     

Metabolic and Physiological Changes Induced by Nitric Oxide and Its Impact on Drought Tolerance in Soybean

Nitric oxide (NO) is an important signaling molecule that plays a pivotal role in stress tolerance. To study the role of NO in drought tolerance and elucidate the underlying mechanisms, NO (0 and 100 μM) was applied to drought-treated soybean plants. Drought stress was imposed by PEG (5% (W/V) of PEG 6000. Nitric oxide improved growth of soybean plants under drought as evidenced by enhanced dry weight (30%). Nitric oxide caused a remarkable increase in activities of catalase and superoxide dismutase (SOD) and SOD expression (14.8-fold), which led to a significant decline in malondealdehyde content under drought conditions. Nitric oxide induced proline biosynthesis due to enhancing pyrroline-5- carboxylate synthetase (P5CS) expression (43.66-fold). The growth-promoting effect of NO application in soybean plants was concomitant with change in metabolic profile (phenolic acid and flavonoid compounds). Nitric oxide up-regulated of phenylalanine ammonia-lyase (PAL) expression in drought-treated plants and may influence on the phenylpropanoid production. Nitric oxide increased salicylic acid (SA) content in soybean plants under stress. So, NO and SA are jointly responsible for boosted tolerance to drought stress in soybean plants. The decrease in unsaturated fatty acid through NO application might reflect a reduction in oxidative damage. These results propose a multifaceted contribution of NO through regulation of physiological and metabolic processes in response to drought stress.

     

Changes in the Phytohormone Levels in Wheat Calli as Affected by Salicylic Acid and Infection with Tilletia caries,a Bunt Pathogenic Agent

The role of salicylic acid (SA) in growth regulation and the change in the levels of phytohormones (IAA, ABA, and cytokinins) were studied in the wheat calli co-cultured with bunt pathogen Tilletia caries. Calli infection with T. caries resulted in the hypertrophied callus growth and simultaneous increase in phytohormone level. The addition of SA to the nutrient media decreased the callus growth induced by the pathogen, whereas the level of investigated phytohormones was not affected. In the SA-treated infected calli, the formation of necrotic lesions was observed in the zones of contact of the fungal mycelium with callus cells that limited pathogen growth. The authors suggest that the stabilization of the hormonal balance of plant cells at pathogenesis is one of the possible mechanisms of the SA protective action in vitro and in vivo. Hence, co-culturing wheat calli and T. caries fungus appeared to be a convenient model for assessing SA protective action.     

Improvement of pistillate flowers yield with GA3 in heavy metals treated plants

GA₃ can be beneficial to improve the flowering yield in Lagenaria siceraria (Mol.) Standl. and Luffa cylindrica L. (Roem) treated with HgCl₂ and . This paper reviews the literature on the possible involvement of gibberellins in flower development and suggests that plants under the stress of heavy metals can be treated with GA₃ to improve growth parameters, to avoid delay in flowering and likewise quality of fruit can be improved. Applied GA₃ (400 mg/l) caused precocious flowering, increasing the number of flowers in the treated plants. All data was compared with control plants grown under similar conditions. Both heavy metals caused significant delay in flowering, consequently leading to reduction in the number of flowers. However, when GA₃ was applied with HgCl₂ and , there was less decrease in staminate and pistillate flowers, revealing the dominant effect of GA₃. Current work reveals that inhibitory effects of heavy metals on flowering can be partially restored with GA₃ application.     

Vesicular arbuscular mycorrhizal mediation of grass response to acidic and heavy metal depositions

Summary The perennial bunchgrassEhrharta calycina was grown with and without V.A.M. fungal infection (Glomus fasciculatum) in a sandy loam exposed to a range of acidic and heavy metal depositions. Heavy metals (Cu, Ni, Pb, Zn, Fe, and Co) were applied in simulated rain (pH 3.0, 4.0, and 5.6) at deposition rates approximating those observed to result from smelter efluents. Metal concentrations in the roots and shoots of mycorrhizal plants were greater than those of non-mycorrhizal plants. Mycorrhizal enhancement of plant metal uptake increased with greater acidity and higher heavy metal content of treatment. The growth of mycorrhizal plants was reduced compared to non-mycorrhizal plants when metal depostion was combined with simulated acid rain. We propose that mycorrhizal enhancement of heavy metal uptake caused reduced growth in plants exposed to acidic and heavy metal depositions.     

Stress Ameliorative Effects of Indole Acetic Acid on <i>Hordeum vulgare</i> L. Seedlings Subjected to Zinc Toxicity

The heavy metals present in the environment accumulate in the plants and affect their productivity and yield. By entering the food chain, metals cause several serious health problems in human beings as well as in other organisms. Indole acetic acid (IAA) is known to act as a signaling molecule between symbiotic association of metal accumulating plants and plant growth promoting rhizobacteria (PGPR). Present study demonstrated a protective role of IAA against surplus Zinc (Zn)-induced toxicity to Hordeum vulgare seedlings. Elevated Zn concentrations suppressed the plant growth, caused a reduction in leaf relative water contents (RWC) and elevated free proline and non-protein thiols (NPT) accumulation. Zinc treatment also led to enhanced lipid peroxidation (MDA contents) as well as the activity of ascorbate peroxidase (APX), showing the involvement of antioxidative defense mechanism to reduce Zn induced toxicity. IAA oxidase activity was also observed to increase due to Zn treatment. IAA pretreatment of H. vulgare caryopsis could partly revert the Zn-induced toxicity in seedlings.     

Salicylic Acid-Mediated Regulation of Morpho-Physiological and Yield Attributes of Wheat and Barley Plants in Deferring Salinity Stress

Salinity has been observed to be a global problem that impede the physiological characteristics of plants. Salicylic acid (SA) as a phytohormone play multifaceted role in plants in terms of development as well as stress management. The current study was conducted to evaluate the effect of salinity and salicylic acid on the performance of wheat and barley plants under field experimentation followed by on-farm study to validate the results. This research was firstly conducted in a 4-year research barley field (2012–2013 and 2013–2014) and wheat (2014–2015 and 2015–2016) and subsequently in an on-farm research in four places (2017–2018). Results depicted that salinity decreased plant yield components and altered ion concentrations (Na⁺/K⁺) causing reduced grain and biological yield. However, SA foliar application induced yield components, especially grain number of plants in both years in non-saline and saline conditions. Exogenously SA application not only led to higher grain yield of barley and wheat but also significantly improved their salt tolerance. Our findings revealed that optimum SA concentrations for achieving highest barley yield were 0.85 and 0.78 mM under saline and non-saline conditions, respectively, while on-farm scale studies observed that foliar application of SA increased grain and biological yield of wheat in Ardakan, Ashkzar (saline soil and water) and Mehrabad (non-saline field) regions. There was no significant effect in Tijerd, a completely non-saline field. The grain yields were higher in SA-treated Ardakan, Ashkzar, and Mehrabad plants in field by 19, 16, and 15%, respectively. Based on present detailed studies, it was concluded that SA improved salinity tolerance and increased crop yield. So, optimum concentration (1.0–1.5 mM) with proper time application (double ridges), SA increased wheat and barley yields up to 20%. Therefore, SA priming could be used as a potent strategy to cope up salinity stress from plants.

     

Salicylic acid induces amelioration of chromium toxicity and affects antioxidant enzyme activity in Sorghum bicolor L.

Aim: Chromium (Cr(VI)) would inflict serious morphological, metabolic, and physiological anomalies in plants ranging from chlorosis of shoot to lipid peroxidation and protein degradation. Cr(VI) toxicity is often associated with oxidative stress, caused by the excessive formation of reactive oxygen species (ROS). In response, plants are equipped with a repertoire of mechanisms to counteract heavy metal (HM) toxicity. Salicylic acid (SA) plays a key role in the signal transduction pathways of various stress responses, demonstrating the protective effect of SA against abiotic stress factors. So, the present investigation was carried out to study the amelioration of pernicious effects of different concentration of Cr(VI) (0.0, 2.0, and 4.0?mg Cr(VI) kg^?1 soil in the form of potassium dichromate) by treatments of salicylic acid solution viz. pretreatment and foliar spray via antioxidative enzymes and their metabolites.

Results: With different treatments of salicylic acid solution, the reinstatement from ill effects of Cr(VI) toxicity was contemplated but the most conspicuous effect was observed when salicylic acid solution was supplied through the foliar spray (0.50?mM). This was accompanied with an increase in ascorbate peroxidase activity and hydrogen peroxide content and decrease in peroxidase activity and ascorbic acid content.

Significance of the study: This study suggests that salicylic acid when applied through pre-treatment of seeds or through a foliar spray can be used to ameliorate the toxic effects of chromium (VI). Salicylic acid has the great potential for reducing the toxicity of heavy metals without negatively impacting the growth of the plants.     

Salicylic and Jasmonic acids in regulation of the proantioxidant state in wheat leaves infected by <Emphasis Type="Italic">Septoria nodorum</Emphasis> Berk

Influence of mediators of the signal systems of salicylic (SA) and jasmonic (JA) acids and their mixture on reactive oxygen species’ (ROS) (superoxide radical and O₂^·− H₂O₂) generation and activity of oxidoreductases (oxalate oxidase, peroxidase and catalase) in leaves of wheat Triticum aestivum L. infected by Septoria leaf blotch pathogen Septoria nodorum Berk has been studied. Presowing treatment of seeds by SA and JA decreased the development rate of fungus on wheat leaves. SA provided earlier inductive effect on production of O₂^·− and H₂O₂ compared with JA. The protective effect of the salicylic and jasmonic acids against Septoria leaf blotch pathogen was caused by activation of oxalate oxidase, induction of anion and cation peroxidases, and decrease of catalase activity. Ability of compounds to stimulate ROS in the plant tissues can be used as criteria for evaluation of immune-modulating activity of new substances for protection of the plants.