Thriving under Salinity: Growth, Ecophysiology and Proteomic Insights into the Tolerance Mechanisms of Obligate Halophyte Suaeda fruticosa.

Studies on obligate halophytes combining eco-physiological techniques and proteomic analysis are crucial for understanding salinity tolerance mechanisms but are limited. We thus examined growth, water relations, ion homeostasis, photosynthesis, oxidative stress mitigation and proteomic responses of an obligate halophyte Suaeda fruticosa to increasing salinity under semi-hydroponic culture. Most biomass parameters increased under moderate (300 mmol L-1 of NaCl) salinity, while high (900 mmol L-1 of NaCl) salinity caused some reduction in biomass parameters. Under moderate salinity, plants showed effective osmotic adjustment with concomitant accumulation of Na+ in both roots and leaves. Accumulation of Na+ did not accompany nutrient deficiency, damage to photosynthetic machinery and oxidative damage in plants treated with 300 mmol L-1 of NaCl. Under high salinity, plants showed further decline in sap osmotic potential with higher Na+ accumulation that did not coincide with a decline in relative water content, Fv/Fm, and oxidative damage markers (H2O2 and MDA). There were 22, 54 and 7 proteins in optimal salinity and 29, 46 and 8 proteins in high salinity treatment that were up-regulated, down-regulated or exhibited no change, respectively, as compared to control plants. These data indicate that biomass reduction in S. fruticosa at high salinity might result primarily from increased energetic cost rather than ionic toxicity.

Bio-inspired synthesis of silver nanoparticles usingSalsola imbricataand its application as antibacterial additive in glass ionomer cement.

Nanotechnology has gained immense popularity and observed rapid development due to the remarkable physio-chemical properties of nanoparticles (NPs) and related nanomaterials. The green production of NPs has many benefits over traditional techniques because the current procedures are expensive, time-consuming, and involve harmful substances that limit their applicability. This study aimed to use a novel green source, theSalsola imbricata(SI) plant, which is commonly found in Central Asia and known for its medicinal properties as a reducing and stabilizing agent for the synthesis of AgNPs. The current study also utilized efficient statistical design, the Plackett-Burman Design (PBD) of Experiment method to synthesize the NPs. The characterization of NPs was carried out using UV-Vis spectroscopy, Fourier transform infrared spectroscopy, and scanning electron microscopy (SEM). The PBD results showed that only two out of four factorsi.e.AgNO3concentration and incubation time, were significant for the synthesis of SI-AgNPs. While remaining factors, incubation temperature and plant extract: AgNO3ratio were non-significant. The SEM analysis result showed that SI-AgNPs had a size of 20-50 nm. The SI-AgNPs demonstrated strong antibacterial activity against oral pathogens such asS. mutans and Lactobacillus acidophilus, with the highest efficacy observed at a concentration of 2 mg ml-1. The addition of SI-AgNPs in glass ionomer cement significantly increased the antibacterial activity of GIC againstS. mutans. Based on the results of the current study, the plant based AgNPs can be further evaluated in detail as alternate antimicrobial agent either alone or in combination with other antimicrobial agents for different dental applications.

The combination of salt and drought benefits selective ion absorption and nutrient use efficiency of halophyte Panicum antidotale.

Soil salinity and water deficit often occur concurrently, but understanding their combined effects on plants' ion regulation is limited. With aim to identify if introducing drought with salinity alleviates salt stress's ionic effects, Panicum antidotale - a halophytic grass- was grown in the presence of single and combined stressors, i.e., drought and salt (low and high). Regulation of cations and anions along with the antioxidant capacity and modifications in leaf anatomy were investigated. Results showed a combination of low salt and drought minimally affected plant (dry) mass by improving the selective ions absorption and nutrient use efficiencies. The lowest ratio for efficiency of photosystem II and carbon assimilation (ΦPSII/ΦCO2) suggested less generation of reactive oxygen species, which were probably detoxified with constitutively performing antioxidant enzymes. In contrast, the combination of high salinity and drought escalated the adverse effects caused due to individual stressors. The selective ion absorption increased, but the non-selective ions transport caused an ionic imbalance indicating the highest ratio of Na+/K+. Although the area of mesophyll increased, a reduction in epidermis (cell number and area) predicted a mechanical injury prone to water loss in these plants. The compromised activity of antioxidant enzymes also suggested treatment-induced oxidative damage. Yet, the synergistic interaction between high salinity and drought was not detrimental to the survival of P. antidotale. Therefore, we suggest planting this grass in habitats with harsh environmental conditions to meet the increasing fodder demands without compromising agricultural lands' productivity.

Biomass Production and Predicted Ethanol Yield Are Linked with Optimum Photosynthesis in Phragmites karka under Salinity and Drought Conditions.

Plant photosynthesis and biomass production are closely associated traits but critical to unfavorable environmental constraints such as salinity and drought. The relationships among stress tolerance, photosynthetic mechanisms, biomass and ethanol yield were assessed in Phragmites karka. The growth parameters, leaf gas exchange and chlorophyll fluorescence of P. karka were studied when irrigated with the control and 100 and 300 mM NaCl in a nutrient solution and water deficit conditions (drought, at 50% water holding capacity). The plant shoot fresh biomass was increased in the low NaCl concentration; however, it significantly declined in high salinity and drought. Interestingly the addition of low salinity increased the shoot biomass and ethanol yield. The number of tillers was increased at 100 mM NaCl in comparison to the control treatment. High salinity increased the photosynthetic performance, but there were no significant changes in drought-treated plants. The saturated irradiance (Is) for photosynthesis increased significantly in low salinity, but it declined (about 50%) in high salt-stressed and (about 20%) in drought-treated plants compared to the control. The rates of dark respiration (Rd) and compensation irradiance (Ic) were decreased significantly under all treatments of salinity and drought, with the exception of unchanged Rd values in the control and drought treatments. A-Ci curve analyses revealed a significant improvement in the Jmax, Vc, max, and triose-phosphate utilization (TPU) at lower salinity levels but decreased at 300 mM NaCl and drought treatments compared to the control. In the chlorophyll fluorescence parameters (Fv/Fm, maximum photochemical quantum yield of PSII, and Y(NO)), the non-photochemical yields were not affected under the salt and drought treatments, although an effective photochemical quantum yield (YII) and electron transport rate (ETR) were significantly enhanced in water deficit compared to control plants. P. karka regulates an efficient photosynthesis mechanism to grow in saline and arid areas and can therefore be used as a sustainable biofuel crop.

Coumarin-Mediated Growth Regulations, Antioxidant Enzyme Activities, and Photosynthetic Efficiency of Sorghum bicolor Under Saline Conditions.

Secondary metabolites, such as phenolic compounds, play an important role in alleviating salinity-induced negative effects in plants. The present study focused on seed priming and foliar application of a potent phenolic compound, coumarin, to induce salinity tolerance in Sorghum bicolor var. SS-77. Based on pilot experiment, 100 mg L-1 concentration of coumarin was applied to mitigate the negative effects of salinity on Sorghum, grown at 0, 100, and 200 mM NaCl under netted greenhouse conditions. Coumarin was applied to each salinity treatment in four different ways (i) non-primed control (NP), (ii) seed priming (COP), (iii) foliar application (COF), and (iv) a combination of seed priming and foliar application (COPF). Salinity stress significantly reduced the plant growth, biochemical attributes, and photosynthetic efficiency of Sorghum, whereas coumarin treatments (COP, COF, and COPF) showed a significant increase (P< 0.01) in above-mentioned parameters at all salinities. Among all, the combined treatment (COPF) showed maximum increase in growth, biochemicals, photosynthetic pigments, antioxidant enzymes, and photosynthetic efficiency parameters. Therefore, it is suggested that a combination of seed priming and foliar spray of 10 mg L-1 coumarin is more suitable than their individual applications. It is an environment friendly and economically feasible approach that will be used to improve salinity tolerance of Sorghum and helpful to get considerable biomass from saline degraded lands to fulfill food, fodder, and energy demands of the ever-growing population.

Combined Transcriptome and Proteome Analysis to Elucidate Salt Tolerance Strategies of the Halophyte Panicum antidotale Retz.

Panicum antidotale, a C4 monocot, has the potential to reclaim saline and drylands and to be utilized as fodder and forage. Its adaptability to survive saline stress has been proven with eco-physiological and biochemical studies. However, little is known about its molecular mechanisms of salt tolerance. In this study, an integrated transcriptome and proteome analysis approach, based on RNA sequencing and liquid chromatography tandem mass spectrometry (LC-MS/MS), was used to identify the said mechanisms. Plants were treated with control (0 mM), low (100 mM), and high (300 mM) sodium chloride (NaCl) treatments to distinguish beneficial and toxic pathways influencing plant biomass. The results indicated differential expression of 3,179 (1,126 upregulated/2,053 downregulated) and 2,172 (898 upregulated/1,274 downregulated) genes (DEGs), and 514 (269 upregulated/245 downregulated) and 836 (494 upregulated/392 downregulated) proteins (DEPs) at 100 and 300 mM NaCl, respectively. Among these, most upregulated genes and proteins were involved in salt resistance strategies such as proline biosynthesis, the antioxidant defense system, ion homeostasis, and sugar accumulation at low salinity levels. On the other hand, the expression of several genes and proteins involved in the respiratory process were downregulated, indicating the inability of plants to meet their energy demands at high salinity levels. Moreover, the impairments in photosynthesis were also evident with the reduced expression of genes regulating the structure of photosystems and increased expression of abscisic acid (ABA) mediated pathways which limits stomatal gas exchange. Similarly, the disturbance in fatty acid metabolism and activation of essential ion transport blockers damaged the integrity of the cell membrane, which was also evident with enhanced malondialdehyde (MDA). Overall, the analysis of pathways revealed that the plant optimal performance at low salinity was related to enhanced metabolism, antioxidative defense, cell growth, and signaling pathways, whereas high salinity inhibited biomass accumulation by altered expression of numerous genes involved in carbon metabolism, signaling, transcription, and translation. The data provided the first global analysis of the mechanisms imparting salt stress tolerance of any halophyte at transcriptome and proteome levels.

Effects of Salinity Stress on Chloroplast Structure and Function.

Salinity is a growing problem affecting soils and agriculture in many parts of the world. The presence of salt in plant cells disrupts many basic metabolic processes, contributing to severe negative effects on plant development and growth. This review focuses on the effects of salinity on chloroplasts, including the structures and function of these organelles. Chloroplasts house various important biochemical reactions, including photosynthesis, most of which are considered essential for plant survival. Salinity can affect these reactions in a number of ways, for example, by changing the chloroplast size, number, lamellar organization, lipid and starch accumulation, and interfering with cross-membrane transportation. Research has shown that maintenance of the normal chloroplast physiology is necessary for the survival of the entire plant. Many plant species have evolved different mechanisms to withstand the harmful effects of salt-induced toxicity on their chloroplasts and its machinery. The differences depend on the plant species and growth stage and can be quite different between salt-sensitive (glycophyte) and salt-tolerant (halophyte) plants. Salt stress tolerance is a complex trait, and many aspects of salt tolerance in plants are not entirely clear yet. In this review, we discuss the different mechanisms of salt stress tolerance in plants with a special focus on chloroplast structure and its functions, including the underlying differences between glycophytes and halophytes.

Synthesis and Characterization of Na-Zeolites from Textile Waste Ash and Its Application for Removal of Lead (Pb) from Wastewater.

Massive production of carcinogenic fly ash waste poses severe threats to water bodies due to its disposal into drains and landfills. Fly ash can be a source of raw materials for the synthesis of adsorbents. Rag fly ash as a new class of raw materials could be a cheap source of Al and Si for the synthesis of Na-zeolites. In this work, NaOH activation, via a prefusion- and postfusion-based hydrothermal strategy, was practiced for the modification of rag fly ash into Na-zeolite. Morphology, surface porosity, chemical composition, functionality, mineral phases, and crystallinity, in conjunction with ion exchangeability of the tailored materials, were evaluated by SEM, ICP-OES, XRF, FTIR, XRD, and cation exchange capacity (CEC) techniques. Rag fly ash and the synthesized Na-zeolites were applied for the removal of Pb (II) from synthetic wastewater by varying the reaction conditions, such as initial metal ion concentration, mass of adsorbent, sorption time, and pH of the reaction medium. It was observed that Na-zeolite materials (1 g/100 mL) effectively removed up to 90-98% of Pb (II) ions from 100 mg/L synthetic solution within 30 min at pH ≈ 8. Freundlich adsorption isotherm favors the multilayer heterogeneous adsorption mechanism for the removal of Pb (II). It is reasonable to conclude that recycling of textile rag fly ash waste into value-added Na-zeolites for the treatment of industrial wastewater could be an emergent move toward achieving sustainable and green remediation.

Salinity induced alterations in photosynthetic and oxidative regulation are ameliorated as a function of salt secretion.

Ion secretion facilitates recretohalophytes to tolerate saline and drought conditions but its relative contribution to the survival of many species remains poorly understood. Tamarix chinensis has high potential for restoration of saline deteriorated lands. The water management and high salt tolerance of the plant have highlighted the need to determine the strategies that govern these mechanisms. Here we report the selectivity of this halophyte to transport, utilize, and secrete different cations and anions under various NaCl (0, 100, 200 and 400 mM) concentrations. Plant growth, photosynthesis and antioxidant defense responses were also determined to relate them with the function of ion secretion. Results reflected two different sets of strategies adopted by plants to survive low and high salinities. Exposure to highly saline conditions caused reduction in photosynthesis due to stomatal and biochemical limitations. The decreased content of photosynthetic pigments exposed plants to excessive light energy that accelerated production of ROS (i.e., hydrogen peroxide H2O2) and caused damage to cellular membranes. The increased activities of anti-oxidative enzymes (superoxide-dismutase, catalase, ascorbate-peroxidase, and glutathione-reductase) were insufficient to detoxify H2O2. In contrast, plants treated with low salinity did not face stomatal limitations while the photosynthetic pigments increased. As no damage to membranes was detected, the increased content of H2O2 was postulated for its messenger role. The assimilation of essential nutrients was affected due to increased content of toxic ions (Na+ and Cl-) in the growing medium and within the plants. However, the ability to regulate K+ facilitated plants to improve water use efficiency under hyper-osmotic environment. The removal of toxic ions from the photosynthesizing tissues demands high energy, which was evident in the compromised growth of plants. This study offers a window to physiological mechanisms, e.g., potassium retention that ensure salt secretion as a beneficial strategy for prolonged survival of T. chinensis.

Trichoderma Enhances Net Photosynthesis, Water Use Efficiency, and Growth of Wheat (Triticum aestivum L.) under Salt Stress.

Soil salinity is one of the most important abiotic stresses limiting plant growth and productivity. The breeding of salt-tolerant wheat cultivars has substantially relieved the adverse effects of salt stress. Complementing these cultivars with growth-promoting microbes has the potential to stimulate and further enhance their salt tolerance. In this study, two fungal isolates, Th4 and Th6, and one bacterial isolate, C7, were isolated. The phylogenetic analyses suggested that these isolates were closely related to Trichoderma yunnanense, Trichoderma afroharzianum, and Bacillus licheniformis, respectively. These isolates produced indole-3-acetic acid (IAA) under salt stress (200 mM). The abilities of these isolates to enhance salt tolerance were investigated by seed coatings on salt-sensitive and salt-tolerant wheat cultivars. Salt stress (S), cultivar (C), and microbial treatment (M) significantly affected water use efficiency. The interaction effect of M x S significantly correlated with all photosynthetic parameters investigated. Treatments with Trichoderma isolates enhanced net photosynthesis, water use efficiency and biomass production. Principal component analysis revealed that the influences of microbial isolates on the photosynthetic parameters of the different wheat cultivars differed substantially. This study illustrated that Trichoderma isolates enhance the growth of wheat under salt stress and demonstrated the potential of using these isolates as plant biostimulants.

Low Salinity Improves Photosynthetic Performance in Panicum antidotale Under Drought Stress.

Salinity and drought are two often simultaneously occurring abiotic stresses that limit the production of food crops worldwide. This study aimed to distinguish between the separate and combined impacts of drought and salinity on the plant response. Panicum antidotale was cultivated in a greenhouse under the following growth conditions: control, 100 mM NaCl (100) and 300 mM NaCl (300) salinity, drought (D; 30% irrigation), and two combinations of salinity and drought (100 + D and 300 + D). The growth response was as follows: 0 ≈ 100 > 100 + D > > D ≈ 300 ≈ 300 + D. Growth correlated directly with photosynthesis. The net photosynthesis, stomatal conductance, intercellular CO2, transpiration, ribulose 1,5-bisphosphate carboxylase (Rubisco), ribulose 1,5-bisphosphate (RuBP) regeneration, and triose phosphate utilization protein (e.g., phosphoenolpyruvate carboxylase) were highest in the control and declined most at 300 + D, while 100 + D performed significantly better as compared to drought. Maximum and actual photosystem II (PSII) efficiencies, along with photochemical quenching during light harvesting, resemble the plant growth and contemporary CO2/H2O gas exchange parameters in the given treatments. Plant improves water use efficiency under salt and drought treatments, which reflects the high water conservation ability of Panicum. Our findings indicate that the combination of low salinity with drought was able to minimize the deleterious effects of drought alone on growth, chlorophyll content, cell integrity, photosynthesis, leaf water potential, and water deficit. This synergetic effect demonstrates the positive role of Na+ and Cl- in carbon assimilation and osmotic adjustment. In contrast, the combination of high salinity and drought enforced the negative response of plants in comparison to single stress, demonstrating the antagonistic impact of water availability and ion toxicity.

Salt induced modulations in antioxidative defense system of Desmostachya bipinnata.

This study addressed the interactions between salt stress and the antioxidant responses of a halophytic grass, Desmostachya bipinnata. Plants were grown in a semi-hydroponic system and treated with different NaCl concentrations (0 mM, 100 mM, 400 mM) for a month. ROS degradation enzyme activities were stimulated by addition of NaCl. Synthesis of antioxidant compounds, such as phenols, was enhanced in the presence of NaCl leading to accumulation of these compounds under moderate salinity. However, when the ROS production rate exceeded the capacity of enzyme-controlled degradation, antioxidant compounds were consumed and oxidative damage was indicated by significant levels of hydrogen peroxide at high salinity. The cellular concentration of salicylic acid increased upon salt stress, but since no direct interaction with ROS was detected, a messenger function may be postulated. High salinity treatment caused a significant decrease of plant growth parameters, whereas treatment with moderate salinity resulted in optimal growth. The activity and abundance of superoxide dismutase (SOD) increased with salinity, but the abundance of SOD isoforms was differentially affected, depending on the NaCl concentration applied. Detoxification of hydrogen peroxide (H2O2) was executed by catalase and guaiacol peroxidase at moderate salinity, whereas the enzymes detoxifying H2O2 through the ascorbate/glutathione cycle dominated at high salinity. The redox status of glutathione was impaired at moderate salinity, whereas the levels of both ascorbate and glutathione significantly decreased only at high salinity. Apparently, the maximal activation of enzyme-controlled ROS degradation was insufficient in comparison to the ROS production at high salinity. As a result, ROS-induced damage could not be prevented, if the applied stress exceeded a critical value in D. bipinnata plants.

Differential protein expression reveals salt tolerance mechanisms of Desmostachya bipinnata at moderate and high levels of salinity.

A proteomics approach was used to investigate salt tolerance mechanisms of Desmostachya bipinnata (L.) Stapf. Plants were subjected to 0mM (control), 100mM (moderate) and 400mM (high) NaCl. Proteins were separated by two-dimensional gel electrophoresis and identified with available databases. Optimal plant fresh weight was found at moderate salinity but declined at high salinity. Water potential, osmotic potential, Na+/K+ ratio, leaf electrolyte leakage, sugars and proline were altered at high salinity. However, water potential, proline content and electrolyte leakage were maintained at moderate salinity; Na+ and K+ concentrations increased, whereas sugars and osmotic potential decreased. Comparative proteome analysis revealed 103 salt responsive proteins. At moderate salinity, most of the proteins involved in energy metabolism, transport, antioxidative defence and cell growth were either unchanged or increased. Proteins related to amino-acid metabolism were decreased while those associated with secondary metabolism were accumulated. At high salinity, amino-acid metabolism and dehydration responses were evident; proteins of energy metabolism, transport and stress defence were downregulated. These results suggest that an efficient defence system, improved transport of water and metabolites, increased cell wall lignification and regulation of energy and carbohydrate metabolism allowed better potential for plant growth under moderately saline conditions.

Effects of salinity and ascorbic acid on growth, water status and antioxidant system in a perennial halophyte.

Salinity causes oxidative stress in plants by enhancing production of reactive oxygen species, so that an efficient antioxidant system, of which ascorbic acid (AsA) is a key component, is an essential requirement of tolerance. However, antioxidant responses of plants to salinity vary considerably among species. Limonium stocksii is a sub-tropical halophyte found in the coastal marshes from Gujarat (India) to Karachi (Pakistan) but little information exists on its salt resistance. In order to investigate the role of AsA in tolerance, 2-month-old plants were treated with 0 (control), 300 (moderate) and 600 (high) mM NaCl for 30 days with or without exogenous application of AsA (20 mM) or distilled water. Shoot growth of unsprayed plants at moderate salinity was similar to that of controls while at high salinity growth was inhibited substantially. Sap osmolality, AsA concentrations and activities of AsA-dependant antioxidant enzymes increased with increasing salinity. Water spray resulted in some improvement in growth, indicating that the growth promotion by exogenous treatments could partly be attributed to water. However, exogenous application of AsA on plants grown under saline conditions improved growth and AsA dependent antioxidant enzymes more than the water control treatment. Our data show that AsA-dependent antioxidant enzymes play an important role in salinity tolerance of L. stocksii.