Auxin Crosstalk with Reactive Oxygen and Nitrogen Species in Plant Development and Abiotic Stress.

The phytohormone auxin acts as an important signaling molecule having regulatory functions during the growth and development of plants. Reactive oxygen species (ROS) are also known to perform signaling functions at low concentrations; however, over-accumulation of ROS due to various environmental stresses damages the biomolecules and cell structures and leads to cell death, and therefore, it can be said that ROS act as a double-edged sword. Nitric oxide (NO), a gaseous signaling molecule, performs a wide range of favorable roles in plants. NO displays its positive role in photomorphogenesis, root growth, leaf expansion, seed germination, stomatal closure, senescence, fruit maturation, mitochondrial activity and metabolism of iron. Studies have revealed the early existence of these crucial molecules during evolution. Moreover, auxin, ROS and NO together show their involvement in various developmental processes and abiotic stress tolerance. Redox signaling is a primary response during exposure of plants to stresses and shows a link with auxin signaling. This review provides updated information related to crosstalk between auxin, ROS and NO starting from their evolution during early Earth periods and their interaction in plant growth and developmental processes as well as in the case of abiotic stresses to plants.

Nitric oxide overcomes copper and copper oxide nanoparticle-induced toxicity in Sorghum vulgare seedlings through regulation of ROS and proline metabolism.

This study aimed to investigate the phytotoxic effect of copper (Cu) and copper nanoparticles (CuONPs) and ameliorative potential of nitric oxide (NO) against these toxic materials in Sorghum vulgare Pers. seedlings. Data suggested that exposure of Cu and CuONPs significantly reduced growth, chlorophyll, carotenoids and protein in root and shoot, which coincided with increased Cu accumulation. However, addition of sodium nitroprusside (SNP, a donor of NO) lowered Cu and CuONPs mediated toxicity through restricting Cu accumulation and improving photosynthetic pigments and total soluble protein contents. Data further suggested that exposure of Cu and CuONPs significantly increased hydrogen peroxide (H2 O2 ), superoxide radicals (O2 •- ), and malondialdehyde (MDA) contents. Enhanced level of oxidative stress severely inhibited the enzymatic activities of glutathione reductase (GR), ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR) and monodehydroascorbate reductase (MDHAR) but enhanced superoxide dismutase (SOD) and catalase (CAT) activity. However, addition of SNP positively regulated antioxidants enzymes activity, particularly the enzymes involved in the ascorbate-glutathione cycle to overcome Cu- and CuONPs-induced stress in Sorghum seedlings. Further, Cu and CuONPs enhanced accumulation of free proline through inducing Δ1 -pyrroline-5-carboxylate synthetase (P5CS) activity while lowering the proline dehydrogenase (PDH) activity. However, addition of SNP reversed these responses. Therefore, overall results revealed that SNP has enough potential of reducing the toxicity of Cu and CuONPs in Sorghum seedlings through regulation of proline metabolism and activity of enzymes of the ascorbate-glutathione cycle. These findings can be employed in developing new resistant varieties of Sorghum having enhanced tolerance against Cu or CuONP stress and improved productivity.

Diverse Physiological Roles of Flavonoids in Plant Environmental Stress Responses and Tolerance.

Flavonoids are characterized as the low molecular weight polyphenolic compounds universally distributed in planta. They are a chemically varied group of secondary metabolites with a broad range of biological activity. The increasing amount of evidence has demonstrated the various physiological functions of flavonoids in stress response. In this paper, we provide a brief introduction to flavonoids' biochemistry and biosynthesis. Then, we review the recent findings on the alternation of flavonoid content under different stress conditions to come up with an overall picture of the mechanism of involvement of flavonoids in plants' response to various abiotic stresses. The participation of flavonoids in antioxidant systems, flavonoid-mediated response to different abiotic stresses, the involvement of flavonoids in stress signaling networks, and the physiological response of plants under stress conditions are discussed in this review. Moreover, molecular and genetic approaches to tailoring flavonoid biosynthesis and regulation under abiotic stress are addressed in this review.

Silicon nanoforms in crop improvement and stress management.

Although, silicon - the second most abundant element in the earth crust could not supersede carbon (C) in the competition of being the building block of life during evolution, yet its presence has been reported in some life forms. In case of the plants, silicon has been reported widely to promote the plant growth under normal as well as stressful situations. Nanoform of silicon is now being explored for its potential to improve plant productivity and its tolerance against various stresses. Silicon nanoparticles (SiNPs) in the form of nanofertilizers, nanoherbicides, nanopesticides, nanosensors and targeted delivery systems, find great utilization in the field of agriculture. However, the mechanisms underlying their uptake by plants need to be deciphered in detail. Silicon nanoformss are reported to enhance plant growth, majorly by improving photosynthesis rate, elevating nutrient uptake and mitigating reactive oxygen species (ROS)-induced oxidative stress. Various studies have reported their ability to provide tolerance against a range of stresses by upregulating plant defense responses. Moreover, they are proclaimed not to have any detrimental impacts on environment yet. This review includes the up-to-date information in context of the eminent role of silicon nanoforms in crop improvement and stress management, supplemented with suggestions for future research in this field.

Metal/Metalloid-Based Nanomaterials for Plant Abiotic Stress Tolerance: An Overview of the Mechanisms.

In agriculture, abiotic stress is one of the critical issues impacting the crop productivity and yield. Such stress factors lead to the generation of reactive oxygen species, membrane damage, and other plant metabolic activities. To neutralize the harmful effects of abiotic stress, several strategies have been employed that include the utilization of nanomaterials. Nanomaterials are now gaining attention worldwide to protect plant growth against abiotic stresses such as drought, salinity, heavy metals, extreme temperatures, flooding, etc. However, their behavior is significantly impacted by the dose in which they are being used in agriculture. Furthermore, the action of nanomaterials in plants under various stresses still require understanding. Hence, with this background, the present review envisages to highlight beneficial role of nanomaterials in plants, their mode of action, and their mechanism in overcoming various abiotic stresses. It also emphasizes upon antioxidant activities of different nanomaterials and their dose-dependent variability in plants' growth under stress. Nevertheless, limitations of using nanomaterials in agriculture are also presented in this review.

Nanoparticles as a potential protective agent for arsenic toxicity alleviation in plants.

Aggrandized technological and industrial progression in past decades have occasioned immense depreciation in the quality of environment and ecosystem, majorly due to augmentation in the number of obnoxious pollutants incessantly being released in soil, water or air. Arsenic (As) is one such hazardous metalloid contaminating the environment which has the potential to detrimentally affect the life on earth. Even in minute quantity, As is known to cause various critical diseases in humans and toxicity in plants. Recent studies on nanoparticles (NPs) approve of their ability to qualify the criterion of becoming a potent tool for mitigating As-induced phytotoxicity. Nanoparticles are reported to promote plant growth under As-stress by stimulating various alterations at physiological, biochemical, and molecular levels. In this review, we provide an up-to-date compilation of research that has been carried out in comprehending the mechanisms utilized by nanoparticles including controlled As uptake and distribution in plants, maintenance of ROS homeostasis during stress and chelation and vacuolar sequestration of As so as to reduce the severity of toxicity induced by As, and potential areas of research in this field will also be indicated for future perspectives.

Silicon and nitric oxide-mediated mechanisms of cadmium toxicity alleviation in wheat seedlings.

The individual impact of silicon (Si) and nitric oxide (NO, as sodium nitroprusside) on metal toxicity in various plant species has been well documented; however, their combined action in the regulation of metal stress has never been tested yet. Therefore, this study investigates the effects of the combined application of Si and NO in the mitigation of Cd toxicity in wheat seedlings. Seedlings grown on Cd has a significantly declined growth due to an increased accumulation of Cd and oxidative stress markers (due to downregulation of antioxidant defense system particularly ascorbate-glutathione cycle) and a decreased accumulation of NO and Si. Additionally, the altered leaf and root structures resulted into a declined photosynthetic efficiency. However, the addition of Si and NO alone as well as combined significantly alleviated Cd toxicity in wheat seedlings by lowering the accumulation of Cd and oxidative stress markers and improving leaf and root structures, which are collectively responsible for a better photosynthetic rate under Cd toxicity, and hence an improved growth was noticed. Particularly, the application of Si and NO in combination lowered the oxidative stress markers via upregulating the antioxidant defense system (particularly AsA-GSH cycle) suggesting the increased efficacy of Si + NO against the Cd toxicity in wheat seedlings as compared to their alone treatments.

Silicon and nitric oxide interplay alleviates copper induced toxicity in mung bean seedlings.

The present study was aimed to investigate copper (Cu) toxicity alleviatory potential of silicon in Vigna radiata L. (mung bean) seedlings. Moreover, attention has also been paid to find out whether endogenous nitric oxide (NO) has any role in Si-governed alleviation of Cu stress. The length of root and shoot, fresh weight, and biochemical attributes were adversely affected by Cu exposure. However, application of Si rescued negative effects of Cu. Cu exposure decreased cell viability, and enhanced cell death and levels of oxidative stress markers (O2•‾, H2O2 and MDA), but Si significantly mitigated these effects of Cu. Application of Cu substantially stimulated the activities of superoxide dismutase and guaiacol peroxidase while inhibited activity of catalase. However, Si addition reversed this effect of Cu. Ascorbate and glutathione contents in roots and shoots were declined by Cu but stimulated by Si. Moreover, we noticed that addition of Nω-nitro-L-arginine methyl ester hydrochloride (L-NAME) and sodium tungstate (Tung) further augmented Cu toxicity but addition of sodium nitroprusside rescued adverse effects of L-NAME and Tung. Altogether, data suggest that though Si was able in alleviating Cu toxicity in mung bean seedlings but it requires endogenous nitric oxide.

A comparative study of the effective response of di-potassium phosphate (K2HPO4) on physiological, biochemical and anatomical aspects of crops dwelling with zinc oxide nanoparticles toxicity.

The dipotassium phosphate (K2HPO4) is a source of phosphorus (P), which is an essential micronutrient for plant growth and reproduction and also acts as a stress alleviator against abiotic stresses. Therefore, it could also become a potential mineral to cope up with zinc oxide nanoparticles' (ZnONPs) toxicity in crops. This study primarily includes synthesis, characterization and differential toxic impacts of ZnONPs on two crop plantsThis study includes synthesis, characterization and differential toxic impacts of ZnONPs on two crop plants, i.e. Triticum aestivum and Solanum lycopersicum, as well as assuage the toxic impacts of ZnONPs through nutrient management approach implied via supplementation of P. The growth and physiological changes under toxic doses of ZnONPs and ameliorative potential of P in crop plants were examined by analysing growth, intracellular Zn accumulation, photosynthetic pigment contents, the kinetics of photosystem II (PS II) photochemistry, root cell anatomy and cell viability via histochemical staining 4',6-diamidino-2-phenylindole and propidium iodide. ZnONPs at 500 and 1000 µM concentrations significantly affected the growth, photosynthetic pigment and PS II photochemistry and cell death in both the plants. It also caused deformation in root anatomy of T. aestivum and S. lycopersicum. Whereas supplementation of P caused significant improvement against ZnONPs stress by causing remarkable enhancement in growth, photosynthetic pigments and activity of PS II photochemistry and decreased cell death. Moreover, the study also discloses the tolerant nature of S. lycopersicum comparing with T. aestivum seedlings. Thus, P is comparatively more effective in managing the ZnONPs toxicity in S. lycopersicum than in T. aestivum.

Aluminum toxicity and aluminum stress-induced physiological tolerance responses in higher plants.

Aluminum (Al) precipitates in acidic soils having a pH < 5.5, in the form of conjugated organic and inorganic ions. Al-containing minerals solubilized in the soil solution cause several negative impacts in plants when taken up along with other nutrients. Moreover, a micromolar concentration of Al present in the soil is enough to induce several irreversible toxicity symptoms such as the rapid and transient over-generation of reactive oxygen species (ROS) such as superoxide anion (O2•-), hydrogen peroxide (H2O2), and hydroxyl radical (•OH), resulting in oxidative bursts. In addition, significant reductions in water and nutrient uptake occur which imposes severe stress in the plants. However, some plants have developed Al-tolerance by stimulating the secretion of organic acids like citrate, malate, and oxalate, from plant roots. Genes responsible for encoding such organic acids, play a critical role in Al tolerance. Several transporters involved in Al resistance mechanisms are members of the Aluminum-activated Malate Transporter (ALMT), Multidrug and Toxic compound Extrusion (MATE), ATP-Binding Cassette (ABC), Natural resistance-associated macrophage protein (Nramp), and aquaporin gene families. Therefore, in the present review, the discussion of the global extension and probable cause of Al in the environment and mechanisms of Al toxicity in plants are followed by detailed emphasis on tolerance mechanisms. We have also identified and categorized the important transporters that secrete organic acids and outlined their role in Al stress tolerance mechanisms in crop plants. The information provided here will be helpful for efficient exploration of the available knowledge to develop Al tolerant crop varieties.

Histochemical Techniques in Plant Science: More Than Meets the Eye.

Histochemistry is an essential analytical tool interfacing extensively with plant science. The literature is indeed constellated with examples showing its use to decipher specific physiological and developmental processes, as well as to study plant cell structures. Plant cell structures are translucent unless they are stained. Histochemistry allows the identification and localization, at the cellular level, of biomolecules and organelles in different types of cells and tissues, based on the use of specific staining reactions and imaging. Histochemical techniques are also widely used for the in vivo localization of promoters in specific tissues, as well as to identify specific cell wall components such as lignin and polysaccharides. Histochemistry also enables the study of plant reactions to environmental constraints, e.g. the production of reactive oxygen species (ROS) can be traced by applying histochemical staining techniques. The possibility of detecting ROS and localizing them at the cellular level is vital in establishing the mechanisms involved in the sensitivity and tolerance to different stress conditions in plants. This review comprehensively highlights the additional value of histochemistry as a complementary technique to high-throughput approaches for the study of the plant response to environmental constraints. Moreover, here we have provided an extensive survey of the available plant histochemical staining methods used for the localization of metals, minerals, secondary metabolites, cell wall components, and the detection of ROS production in plant cells. The use of recent technological advances like CRISPR/Cas9-based genome-editing for histological application is also addressed. This review also surveys the available literature data on histochemical techniques used to study the response of plants to abiotic stresses and to identify the effects at the tissue and cell levels.

Regulation of ascorbate-glutathione cycle by exogenous nitric oxide and hydrogen peroxide in soybean roots under arsenate stress.

The role of nitric oxide (NO) and hydrogen peroxide (H2O2) is well known for regulating plant abiotic stress responses. However, underlying mechanisms are still poorly understood. Therefore, the present study investigated the involvement of NO and H2O2 signalling in the regulation of arsenate toxicity (AsV) in soybean roots employing a pharmacological approach. Results show that AsV toxicity declined root length and biomass due to greater As accumulation in the cell wall and cellular organelles. Arsenate induced cell death due to enhanced levels of reactive oxygen species, lipid and protein oxidation and down-regulation in ascorbate-glutathione cycle and redox states of ascorbate and glutathione. These results correlate with lower endogenous level of NO. Interestingly, addition of L-NAME increased AsV toxicity. However, addition of SNP reverses effect of L-NAME, suggesting that endogenous NO has a role in mitigating AsV toxicity. Exogenous H2O2 also demonstrated capability of alleviating AsV stress, while NAC reversed the protective effect of H2O2. Furthermore, DPI application further increased AsV toxicity, suggesting that endogenous H2O2 is also implicated in mitigating AsV stress. SNP was not able to mitigate AsV toxicity in the presence of DPI, suggesting that H2O2 might have acted downstream of NO in accomplishing amelioration of AsV toxicity.

Structural modifications of plant organs and tissues by metals and metalloids in the environment: A review.

At the dawn of the industrial revolution, the exorbitant use of heavy metals and toxic elements by mankind unfurls a powerful and complex web of hazard all around the world that significantly contributed to unprecedented trends in environmental degradation. Plants as sessile organisms, that cannot escape from the stress directly, have adapted to this environment via concurrent configurations of several traits. Among them the anatomy has been identified as much more advanced field of research that brought the explosion of interest among the expertise and its prodigious importance in stress physiology is unavoidable. In conjunction with various other disciplines, like physiology, biochemistry, genomics and metabolomics, the plant anatomy provides a large data sets that are paving the way towards a comprehensive and holistic understanding of plant growth, development, defense and productivity under heavy metal and toxic element stress. Present paper advances our recent knowledge about structural alterations of plant tissues induced by metals and metalloids, like antimony (Sb), arsenic (As), aluminium (Al), copper (Cu), cadmium (Cd), chromium (Cr), lead (Pb), manganese (Mn), mercury (Hg), nickel (Ni) and zinc (Zn) and points on essential role of plant anatomy and its understanding for plant growth and development in changing environment. Understanding of anatomical adaptations of various plant organs and tissues to heavy metals and metalloids could greatly contribute to integral and modern approach for investigation of plants in changing environmental conditions. These findings are necessary for understanding of the whole spectra of physiological and biochemical reactions in plants and to maintain the crop productivity worldwide. Moreover, our holistic perception regarding the processes underlying the plant responses to metal(loids) at anatomical level are needed for improving crop management and breeding techniques.

Metabolic Syndrome and Its Effects on Cartilage Degeneration vs Regeneration: A Pilot Study Using Osteoarthritis Biomarkers.

BACKGROUND: Osteoarthritis (OA) of the knee is one of the leading causes of disability characterized by degeneration of hyaline cartilage combined with reparative processes. Its strong association with metabolic syndrome is postulated to be due to both mechanical and biochemical factors. Our study aims to study differential effect of metabolic risk factors on cartilage degeneration and regeneration at biomarker level. DESIGN: After screening 281 patients presenting with knee pain, 41 patients who met the selection criteria were included and were divided into metabolic (MetS) OA and non-metabolic (Non-MetS) OA phenotypes using National Cholesterol Education Programme-Adult Treatment Panel-III (NCEP-ATP-III) criteria for metabolic syndrome. Serum Cartilage Oligomeric Matrix Protein (COMP) and Procollagen type IIA N terminal Propeptide (PIIANP) levels were used as tools to assess cartilage degeneration and regeneration, respectively. RESULTS: 22 among 41 patients (53.66%) had metabolic syndrome. Covariates like age, gender, Kellgren Lawrence (KL) grades were comparable in both groups. MetS-OA group showed significant increase in serum COMP levels (p = 0.03) with no significant effect on serum PIIANP levels (p = 0.46). Hypertriglyceridemia showed independent association with both cartilage anabolism (p = 0.03) and catabolism (p = 0.03). CONCLUSION: Metabolic syndrome, though has no effect on cartilage regeneration tends to shift cartilage homeostasis towards degeneration with hypertriglyceridemia showing significant independent effect on cartilage metabolism.

Silicon tackles butachlor toxicity in rice seedlings by regulating anatomical characteristics, ascorbate-glutathione cycle, proline metabolism and levels of nutrients.

Reckless use of herbicides like butachlor (Buta) in the fields represents a serious threat to crop plants, and hence to their productivity. Silicon (Si) is well known for its implication in the alleviation of the effects of abiotic stresses; however, its role in mitigating Buta toxicity is not yet known. Therefore, this study was carried out to explore the role of Si (10 µM) in regulating Buta (4 µM) toxicity in rice seedlings. Buta reduced growth and photosynthesis, altered nitric oxide (NO) level and leaf and root anatomy, inhibited enzyme activities of the ascorbate-glutathione cycle (while transcripts of associated enzymes, increased except OsMDHAR), as well as its metabolites (ascorbate and glutathione) and uptake of nutrients (Mg, P, K, S, Ca, Fe, etc. except Na), while addition of Si reversed Buta-induced alterations. Buta stimulated the expression of Si channel and efflux transporter genes- Lsi1 and Lsi2 while the addition of Si further greatly induced their expression under Buta toxicity. Buta increased free proline accumulation by inducing the activity of Δ1-pyrroline-5-carboxylate synthetase (P5CS) and decreasing proline dehydrogenase (PDH) activity, while Si reversed these effects caused by Buta. Our results suggest that Si-governed mitigation of Buta toxicity is linked with favorable modifications in energy flux parameters of photosynthesis and leaf and root anatomy, up-regulation of Si channel and transporter genes, ascorbate-glutathione cycle and nutrient uptake, and lowering in oxidative stress. We additionally demonstrate that NO might have a crucial role in these responses.

Fascinating impact of silicon and silicon transporters in plants: A review.

Silicon (Si) is a metalloid which is gaining worldwide attention of plant scientists due to its ameliorating impact on plants' growth and development. The beneficial response of Si is observed predominantly under numerous abiotic and biotic stress conditions. However, under favorable conditions, most of the plant can grow without it. Therefore, Si has yet not been fully accepted as essential element rather it is being considered as quasi-essential for plants' growth. Si is also known to enhance resilience in plants by reducing the plant's stress. Besides its second most abundance on the earth crust, most of the soils lack plant available form of Si i.e. silicic acid. In this regard, understanding the role of Si in plant metabolism, its uptake from roots and transport to aerial tissues along with its ionomics and proteomics under different circumstances is of great concern. Plants have evolved a well-optimized Si-transport system including various transporter proteins like Low silicon1 (Lsi1), Low silicon2 (Lsi2), Low silicon3 (Lsi3) and Low silicon6 (Lsi6) at specific sub-cellular locations along with the expression profiling that creates precisely coordinated network among these transporters, which also facilitate uptake and accumulation of Si. Though, an ample amount of information is available pertinent to the solute specificity, active sites, transcriptional and post-transcriptional regulation of these transporter genes. Similarly, the information regarding transporters involved in Si accumulation in different organelles is also available particularly in silica cells occurred in poales. But in this review, we have attempted to compile studies related to plants vis à vis Si, its role in abiotic and biotic stress, its uptake in various parts of plants via different types of Si-transporters, expression pattern, localization and the solute specificity. Besides these, this review will also provide the compiled knowledge about the genetic variation among crop plants vis à vis enhanced Si uptake and related benefits.