Regulation of ROS Metabolism in Plants under Environmental Stress: A Review of Recent Experimental Evidence.

Various environmental stresses singly or in combination generate excess amounts of reactive oxygen species (ROS), leading to oxidative stress and impaired redox homeostasis. Generation of ROS is the obvious outcome of abiotic stresses and is gaining importance not only for their ubiquitous generation and subsequent damaging effects in plants but also for their diversified roles in signaling cascade, affecting other biomolecules, hormones concerning growth, development, or regulation of stress tolerance. Therefore, a good balance between ROS generation and the antioxidant defense system protects photosynthetic machinery, maintains membrane integrity, and prevents damage to nucleic acids and proteins. Notably, the antioxidant defense system not only scavenges ROS but also regulates the ROS titer for signaling. A glut of studies have been executed over the last few decades to discover the pattern of ROS generation and ROS scavenging. Reports suggested a sharp threshold level of ROS for being beneficial or toxic, depending on the plant species, their growth stages, types of abiotic stresses, stress intensity, and duration. Approaches towards enhancing the antioxidant defense in plants is one of the vital areas of research for plant biologists. Therefore, in this review, we accumulated and discussed the physicochemical basis of ROS production, cellular compartment-specific ROS generation pathways, and their possible distressing effects. Moreover, the function of the antioxidant defense system for detoxification and homeostasis of ROS for maximizing defense is also discussed in light of the latest research endeavors and experimental evidence.

Glutathione in plants: biosynthesis and physiological role in environmental stress tolerance.

Glutathione (GSH; γ-glutamyl-cysteinyl-glycine) is a small intracellular thiol molecule which is considered as a strong non-enzymatic antioxidant. Glutathione regulates multiple metabolic functions; for example, it protects membranes by maintaining the reduced state of both α-tocopherol and zeaxanthin, it prevents the oxidative denaturation of proteins under stress conditions by protecting their thiol groups, and it serves as a substrate for both glutathione peroxidase and glutathione S-transferase. By acting as a precursor of phytochelatins, GSH helps in the chelating of toxic metals/metalloids which are then transported and sequestered in the vacuole. The glyoxalase pathway (consisting of glyoxalase I and glyoxalase II enzymes) for detoxification of methylglyoxal, a cytotoxic molecule, also requires GSH in the first reaction step. For these reasons, much attention has recently been directed to elucidation of the role of this molecule in conferring tolerance to abiotic stress. Recently, this molecule has drawn much attention because of its interaction with other signaling molecules and phytohormones. In this review, we have discussed the recent progress in GSH biosynthesis, metabolism and its role in abiotic stress tolerance.

Attenuation of negative effects caused by a combination of heat and cadmium stress in Arabidopsis thaliana deficient in jasmonic acid synthesis.

Due to recent global warming, heat stress can simultaneously occur with cadmium (Cd) stress in regions suffering from metal pollution. In this study, we investigated the effects of heat, Cd and their combination on the growth and physiological characteristics of Arabidopsis thaliana. Arabidopsis plants were more susceptible to a combination of heat and Cd stress than to each stress applied individually, although the accumulation of Cd in shoots was comparable between plants subjected to Cd stress and the combined stress. Plants subjected to this stress combination showed a dramatic reduction in the accumulation of the photosynthetic reaction center proteins in photosystem II as well as a tendency toward enhanced lipid peroxidation, suggesting that the negative effects of a combination of heat and Cd stresses might be caused by oxidative damage accompanied by damage to the photosynthetic apparatus. Interestingly, aos and lox3 mutants deficient in jasmonic acid (JA) synthesis showed attenuation of the negative effects caused by a combination of heat and Cd stresses on the growth and maximum quantum efficiency of photosystem II. The roles of JA might be altered when heat stress is combined with Cd stress, despite its significance in the tolerance of plants to Cd stress when individually applied, which has been shown in previous studies.

Biostimulants for the Regulation of Reactive Oxygen Species Metabolism in Plants under Abiotic Stress.

Global food security for a growing population with finite resources is often challenged by multiple, simultaneously occurring on-farm abiotic stresses (i.e., drought, salinity, low and high temperature, waterlogging, metal toxicity, etc.) due to climatic uncertainties and variability. Breeding for multiple stress tolerance is a long-term solution, though developing multiple-stress-tolerant crop varieties is still a challenge. Generation of reactive oxygen species in plant cells is a common response under diverse multiple abiotic stresses which play dual role of signaling molecules or damaging agents depending on concentration. Thus, a delicate balance of reactive oxygen species generation under stress may improve crop health, which depends on the natural antioxidant defense system of the plants. Biostimulants represent a promising type of environment-friendly formulation based on natural products that are frequently used exogenously to enhance abiotic stress tolerance. In this review, we illustrate the potential of diverse biostimulants on the activity of the antioxidant defense system of major crop plants under stress conditions and their other roles in the management of abiotic stresses. Biostimulants have the potential to overcome oxidative stress, though their wider applicability is tightly regulated by dose, crop growth stage, variety and type of biostimulants. However, these limitations can be overcome with the understanding of biostimulants' interaction with ROS signaling and the antioxidant defense system of the plants.

Oxidative Damage and Antioxidant Defense in Sesamum indicum after Different Waterlogging Durations.

The present study was designed to investigate the duration-dependent changes in the biochemical attributes of sesame in response to waterlogging stress. Sesame plants (Sesamum indicum L. cv. BARI Til-4) were subjected to waterlogging for 2, 4, 6, and 8 days during the vegetative stage and data were measured following waterlogging treatment. The present study proves that waterlogging causes severe damage to different attributes of the sesame plant. The plants showed an increasing trend in lipid peroxidation as well as hydrogen peroxide (H2O2) and methylglyoxal contents that corresponded to increased stress duration. A prolonged period of waterlogging decreased leaf relative water content and proline content. Photosynthetic pigments, like chlorophyll (chl) a, b, and chl (a+b) and carotenoid contents, also decreased over time in stressed plants. Glutathione (GSH) and oxidized glutathione (GSSG) contents increased under waterlogging, while the GSH/GSSG ratio and ascorbate content decreased, indicating the disruption of redox balance in the cell. Ascorbate peroxidase, monodehydroascorbate reductase, and glutathione peroxidase activity increased under waterlogging, while dehydroascorbate reductase, glutathione reductase, and catalase activity mostly decreased. Waterlogging modulated the glyoxalase system mostly by enhancing glyoxalase II activity, with a slight increase in glyoxalase I activity. The present study also demonstrates the induction of oxidative stress via waterlogging in sesame plants and that stress levels increase with increased waterlogging duration.

Regulation of Ascorbate-Glutathione Pathway in Mitigating Oxidative Damage in Plants under Abiotic Stress.

Reactive oxygen species (ROS) generation is a usual phenomenon in a plant both under a normal and stressed condition. However, under unfavorable or adverse conditions, ROS production exceeds the capacity of the antioxidant defense system. Both non-enzymatic and enzymatic components of the antioxidant defense system either detoxify or scavenge ROS and mitigate their deleterious effects. The Ascorbate-Glutathione (AsA-GSH) pathway, also known as Asada-Halliwell pathway comprises of AsA, GSH, and four enzymes viz. ascorbate peroxidase, monodehydroascorbate reductase, dehydroascorbate reductase, and glutathione reductase, play a vital role in detoxifying ROS. Apart from ROS detoxification, they also interact with other defense systems in plants and protect the plants from various abiotic stress-induced damages. Several plant studies revealed that the upregulation or overexpression of AsA-GSH pathway enzymes and the enhancement of the AsA and GSH levels conferred plants better tolerance to abiotic stresses by reducing the ROS. In this review, we summarize the recent progress of the research on AsA-GSH pathway in terms of oxidative stress tolerance in plants. We also focus on the defense mechanisms as well as molecular interactions.

Exogenous Silicon Attenuates Cadmium-Induced Oxidative Stress in Brassica napus L. by Modulating AsA-GSH Pathway and Glyoxalase System.

Cadmium (Cd) brings a devastating health hazard to human being as a serious consequence of agricultural and environmental contamination. We demonstrated the protective effect of silicon (Si) on cadmium (Cd)-stressed rapeseed (Brassica napus L. cv. BINA Sharisha 3) plants through regulation of antioxidant defense and glyoxalase systems. Twelve-day-old seedlings were exposed to Cd stress (0.5 and 1.0 mM CdCl2) separately and in combination with Si (SiO2, 1.0 mM) for 2 days. Cadmium toxicity was evident by an obvious oxidative stress through sharp increases in H2O2 content and lipid peroxidation (malondialdehyde, MDA content), and visible sign of superoxide and H2O2. Cadmium stress also decreased the content of ascorbate (AsA) and glutathione (GSH) as well as their redox pool. The activities of monodehydroascorbate reductase (MDHAR), dehydroascorbate reductase (DHAR) and catalase (CAT) were decreased by Cd while ascorbate peroxidase (APX) and glutathione S-transferase (GST) activities were increased. The enzymes of glyoxalase system (glyoxalase I, Gly I and glyoxalase II, Gly II) were also inefficient under Cd stress. However, exogenous application of Si in Cd treated seedlings reduced H2O2 and MDA contents and improved antioxidant defense mechanism through increasing the AsA and GSH pools and activities of AsA-GSH cycle (APX, MDHAR, DHAR and GR) and glyoxalase system (Gly I and Gly II) enzymes and CAT. Thus Si reduced oxidative damage in plants to make more tolerant under Cd stress through augmentation of different antioxidant components and methylglyoxal detoxification system.