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Journal ArticleDOI

Reactive oxygen species (ROS) and response of antioxidants as ROS-scavengers during environmental stress in plants

TL;DR: This review focuses on the different types of ROS, their cellular production sites, their targets, and their scavenging mechanism mediated by both the branches of the antioxidant systems, highlighting the potential role of antioxidant in plants.
Abstract: Reactive oxygen species (ROS) were initially recognized as toxic by-products of aerobic metabolism. In recent years, it has become apparent that ROS plays an important signaling role in plants, controlling processes such as growth, development and especially response to biotic and abiotic environmental stimuli. The major members of the ROS family include free radicals like O2● −, OH● and non-radicals like H2O2 and 1O2. The ROS production in plants is mainly localized in the chloroplast, mitochondria and peroxisomes. There are secondary sites as well like the endoplasmic reticulum, cell membrane, cell wall and the apoplast. The role of the ROS family is that of a double edged sword; while they act as secondary messengers in various key physiological phenomena, they also induce oxidative damages under several environmental stress conditions like salinity, drought, cold, heavy metals, UV irradiation etc., when the delicate balance between ROS production and elimination, necessary for normal cellular homeostasis, is disturbed. The cellular damages are manifested in the form of degradation of biomolecules like pigments, proteins, lipids, carbohydrates and DNA, which ultimately amalgamate in plant cellular death. To ensure survival, plants have developed efficient antioxidant machinery having two arms, (i) enzymatic components like superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX), guaiacol peroxidase (GPX), glutathione reductase (GR), monodehydroascorbate reductase (MDHAR) and dehydroascorbate reductase (DHAR); (ii) non-enzymatic antioxidants like ascorbic acid (AA), reduced glutathione (GSH), α-tocopherol, carotenoids, flavonoids and the osmolyte proline. These two components work hand in hand to scavenge ROS. In this review, we emphasize on the different types of ROS, their cellular production sites, their targets, and their scavenging mechanism mediated by both the branches of the antioxidant systems, highlighting the potential role of antioxidant
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Journal ArticleDOI
TL;DR: A need to understand the processes and role of oxidative stress in neurodegenerative diseases is understood, with a focus on the pivotal role played by OS in mitochondrial dysfunction.
Abstract: Oxidative stress is proposed as a regulatory element in ageing and various neurological disorders. The excess of oxidants causes a reduction of antioxidants, which in turn produce an oxidation–reduction imbalance in organisms. Paucity of the antioxidant system generates oxidative-stress, characterized by elevated levels of reactive species (oxygen, hydroxyl free radical, and so on). Mitochondria play a key role in ATP supply to cells via oxidative phosphorylation, as well as synthesis of essential biological molecules. Various redox reactions catalyzed by enzymes take place in the oxidative phosphorylation process. An inefficient oxidative phosphorylation may generate reactive oxygen species (ROS), leading to mitochondrial dysfunction. Mitochondrial redox metabolism, phospholipid metabolism, and proteolytic pathways are found to be the major and potential source of free radicals. A lower concentration of ROS is essential for normal cellular signaling, whereas the higher concentration and long-time exposure of ROS cause damage to cellular macromolecules such as DNA, lipids and proteins, ultimately resulting in necrosis and apoptotic cell death. Normal and proper functioning of the central nervous system (CNS) is entirely dependent on the chemical integrity of brain. It is well established that the brain consumes a large amount of oxygen and is highly rich in lipid content, becoming prone to oxidative stress. A high consumption of oxygen leads to excessive production of ROS. Apart from this, the neuronal membranes are found to be rich in polyunsaturated fatty acids, which are highly susceptible to ROS. Various neurodegenerative diseases such as Parkinson’s disease (PD), Alzheimer’s disease (AD), Huntington’s disease (HD), and amyotrophic lateral sclerosis (ALS), among others, can be the result of biochemical alteration (due to oxidative stress) in bimolecular components. There is a need to understand the processes and role of oxidative stress in neurodegenerative diseases. This review is an effort towards improving our understanding of the pivotal role played by OS in neurodegenerative disorders.

920 citations

Journal ArticleDOI
TL;DR: This review describes recent progress in deciphering the mechanisms controlling ion homeostasis, cell activity responses, and epigenetic regulation in plants under salt stress and highlights research areas that require further research to reveal new determinants of salt tolerance in plants.
Abstract: Contents Summary 523 I. Introduction 523 II. Sensing salt stress 524 III. Ion homeostasis regulation 524 IV. Metabolite and cell activity responses to salt stress 527 V. Conclusions and perspectives 532 Acknowledgements 533 References 533 SUMMARY: Excess soluble salts in soil (saline soils) are harmful to most plants. Salt imposes osmotic, ionic, and secondary stresses on plants. Over the past two decades, many determinants of salt tolerance and their regulatory mechanisms have been identified and characterized using molecular genetics and genomics approaches. This review describes recent progress in deciphering the mechanisms controlling ion homeostasis, cell activity responses, and epigenetic regulation in plants under salt stress. Finally, we highlight research areas that require further research to reveal new determinants of salt tolerance in plants.

703 citations


Cites background from "Reactive oxygen species (ROS) and r..."

  • ...2AA +NAD Vitis vinifera,Oryza sativa Seedling Reviewed by Das & Roychoudhury (2014); Ikbal et al. (2014); Mostofa et al. (2015); Rahman et al. (2016) GR GSSG +NADPH?...

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  • ...…reductase (DHAR), monodehydroascorbate reductase (MDHAR), glutathione peroxidase (GR), and glutathione S-transferase (GST) (Sreenivasulu et al., 2000; Meloni et al., 2003; Amako & Ushimaru, 2009; Das & Roychoudhury, 2014; Begaramorales et al., 2015; Li et al., 2015; Wang et al., 2017)....

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  • ...2GSH +NADP+ Echinochloa crusgalli, Brassica juncea,Oryza sativa Seedling, leaf, root Abogadallah et al. (2010); reviewed by Das & Roychoudhury (2014);Mostofa et al. (2015); Ranjit et al. (2016) SOD, superoxide dismutase; APX, ascorbate peroxidase; CAT, catalase; GPX, guaiacolperoxidase; DHAR, dehydroascorbate reductase; MDHAR, monodehydroascorbate reductase; GR, glutathione peroxidase; AA, ascorbic acid; DHA, reduces dehydroascorbate; GSSG, glutathione dissulfide; GSH, reduced glutathione; MDHA, monodehydroascorbate; NADH, nicotinamide adenine dinucleotide; NAD, nicotinamide adenine dinucleotide; NADPH, nicotinamide adenine dinucleotide phosphate; NADP+, oxidized form of nicotinamide adenine dinucleotide phosphate....

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  • ...Overexpression of an enzyme either removing ROS or accumulating ROS scavengers may increase plant salt tolerance (Das & Roychoudhury, 2014)....

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  • ...MT reorganization is regulated by the ras homologus oncogenes (Rho)related GTPase from plants (ROP2) and the ROP-interactive GPX H2O2 +DHA?...

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Journal ArticleDOI
TL;DR: Current knowledge of the possible mechanisms associated with H2O2-induced abiotic oxidative stress tolerance in plants is reviewed, with special reference to antioxidant metabolism.
Abstract: Plants are constantly challenged by various abiotic stresses that negatively affect growth and productivity worldwide. During the course of their evolution, plants have developed sophisticated mechanisms to recognize external signals allowing them to respond appropriately to environmental conditions, although the degree of adjustability or tolerance to specific stresses differs from species to species. Overproduction of reactive oxygen species (ROS) (hydrogen peroxide, H2O2; superoxide, O2ˉ˙; hydroxyl radical, OH. and singlet oxygen, 1O2) is enhanced under abiotic and/or biotic stresses, which can cause oxidative damage to plant macromolecules and cell structures, leading to inhibition of plant growth and development, or to death. Among the various ROS, freely diffusible and relatively long-lived H2O2 acts as a central player in stress signal transduction pathways. These pathways can then activate multiple acclamatory responses that reinforce resistance to various abiotic and biotic stressors. To utilize H2O2 as a signaling molecule, non-toxic levels must be maintained in a delicate balancing act between H2O2 production and scavenging. Several recent studies have demonstrated that the H2O2-priming can enhance abiotic stress tolerance by modulating ROS detoxification and by regulating multiple stress-responsive pathways and gene expression. Despite the importance of the H2O2-priming, little is known about how this process improves the tolerance of plants to stress. Understanding the mechanisms of H2O2-priming-induced abiotic stress tolerance will be valuable for identifying biotechnological strategies to improve abiotic stress tolerance in crop plants. This review is an overview of our current knowledge of the possible mechanisms associated with H2O2-induced abiotic oxidative stress tolerance in plants, with special reference to antioxidant metabolism.

553 citations


Cites background from "Reactive oxygen species (ROS) and r..."

  • ...…), the hydroxyl radical (OH·) and singlet oxygen (1O·2) are also produced as one of the earliest responses of plant cells to environmental stresses, and these ROS molecules can cause damage to a variety of biological processes (Halliwell, 2006; Gill and Tuteja, 2010; Das and Roychoudhury, 2014)....

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  • ...…and metal or metalloid stresses, ROS levels can rise significantly, leading to redox imbalance and oxidative stress (Hossain et al., 2010; Hasanuzzaman et al., 2011a,b; Hossain and Fujita, 2013; Mostofa and Fujita, 2013; Das and Roychoudhury, 2014; Mostofa et al., 2014a,b,c; Nahar et al., 2014)....

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Journal ArticleDOI
TL;DR: This review highlights the importance of the As-induced generation of reactive oxygen species (ROS) as well as their damaging impacts on plants at biochemical, genetic, and molecular levels.
Abstract: Environmental contamination with arsenic (As) is a global environmental, agricultural and health issue due to the highly toxic and carcinogenic nature of As. Exposure of plants to As, even at very low concentration, can cause many morphological, physiological, and biochemical changes. The recent research on As in the soil-plant system indicates that As toxicity to plants varies with its speciation in plants (e.g., arsenite, As(III); arsenate, As(V)), with the type of plant species, and with other soil factors controlling As accumulation in plants. Various plant species have different mechanisms of As(III) or As(V) uptake, toxicity, and detoxification. This review briefly describes the sources and global extent of As contamination and As speciation in soil. We discuss different mechanisms responsible for As(III) and As(V) uptake, toxicity, and detoxification in plants, at physiological, biochemical, and molecular levels. This review highlights the importance of the As-induced generation of reactive oxygen species (ROS), as well as their damaging impacts on plants at biochemical, genetic, and molecular levels. The role of different enzymatic (superoxide dismutase, catalase, glutathione reductase, and ascorbate peroxidase) and non-enzymatic (salicylic acid, proline, phytochelatins, glutathione, nitric oxide, and phosphorous) substances under As(III/V) stress have been delineated via conceptual models showing As translocation and toxicity pathways in plant species. Significantly, this review addresses the current, albeit partially understood, emerging aspects on (i) As-induced physiological, biochemical, and genotoxic mechanisms and responses in plants and (ii) the roles of different molecules in modulation of As-induced toxicities in plants. We also provide insight on some important research gaps that need to be filled to advance our scientific understanding in this area of research on As in soil-plant systems.

513 citations


Cites background from "Reactive oxygen species (ROS) and r..."

  • ...Due to increased peroxidation of PUFA, there is a decrease in membrane fluidity and increases in leakiness, causing severe damage to membrane proteins [228]....

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  • ...Aldehydes formed in mitochondria are demonstrated to impose cytoplasmic male sterility in Zea mays [228]....

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Journal ArticleDOI
TL;DR: Over the past two decades it has been proven that ROS together with non‐photochemical quenching (NPQ), hormones, Ca2+ waves, and electrical signals are the main players in SAA and SAR, two physiological processes essential for plant survival and productivity in unfavorable conditions.

377 citations


Cites background from "Reactive oxygen species (ROS) and r..."

  • ...animals, in plant cells ROS are generated in different subcellular compartments, mainly in the chloroplasts and peroxisomes, but also in the mitochondria, plasma membrane, cell wall, endoplasmic reticulum and nuclei [10,11]....

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  • ...for photosystem II activity loss, which can also lead to plant cell death [10]....

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  • ...scavenging this toxic radical, its excess accumulation triggers programmed cell death [10]....

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References
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Journal ArticleDOI
TL;DR: The mechanisms of ROS generation and removal in plants during development and under biotic and abiotic stress conditions are described and the possible functions and mechanisms for ROS sensing and signaling in plants are compared with those in animals and yeast.
Abstract: Several reactive oxygen species (ROS) are continuously produced in plants as byproducts of aerobic metabolism. Depending on the nature of the ROS species, some are highly toxic and rapidly detoxified by various cellular enzymatic and nonenzymatic mechanisms. Whereas plants are surfeited with mechanisms to combat increased ROS levels during abiotic stress conditions, in other circumstances plants appear to purposefully generate ROS as signaling molecules to control various processes including pathogen defense, programmed cell death, and stomatal behavior. This review describes the mechanisms of ROS generation and removal in plants during development and under biotic and abiotic stress conditions. New insights into the complexity and roles that ROS play in plants have come from genetic analyses of ROS detoxifying and signaling mutants. Considering recent ROS-induced genome-wide expression analyses, the possible functions and mechanisms for ROS sensing and signaling in plants are compared with those in animals and yeast.

9,908 citations


"Reactive oxygen species (ROS) and r..." refers background in this paper

  • ...These are very lethal and causes extensive damage to protein, DNA and lipids and thereby affects normal cellular functioning (Apel and Hirt, 2004; Foyer and Noctor, 2005)....

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  • ...The NADPH-dependent-oxidases which are localized in the plasma membrane are in the spotlight due to their gene expression and presence of different homologs during different stress conditions (Apel and Hirt, 2004)....

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Journal ArticleDOI
TL;DR: Key steps of the signal transduction pathway that senses ROIs in plants have been identified and raise several intriguing questions about the relationships between ROI signaling, ROI stress and the production and scavenging ofROIs in the different cellular compartments.

9,395 citations


"Reactive oxygen species (ROS) and r..." refers background in this paper

  • ...Since it regenerates AA, it is co-localized with the APX in the peroxisomes and mitochondria, where APX scavenges H2O2 and oxidizes AA in the process (Mittler, 2002)....

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  • ...SODs are classified into three isozymes based on the metal ion it binds, Mn-SOD (localized in mitochondria), Fe-SOD (localized in chloroplasts), and Cu/Zn-SOD (localized in cytosol, peroxisomes, and chloroplasts) (Mittler, 2002)....

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  • ...The NADPH-mediated electron transport involving CytP450, localized in the ER generates O•−2 (Mittler, 2002)....

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  • ...Peroxisomes are the hotspots of H2O2 production due to β-oxidation of fatty acids, photorespiration, purine catabolism and oxidative stress (Mittler, 2002)....

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Journal ArticleDOI
TL;DR: The biochemistry of ROS and their production sites, and ROS scavenging antioxidant defense machinery are described, which protects plants against oxidative stress damages.

8,259 citations


"Reactive oxygen species (ROS) and r..." refers background in this paper

  • ...…peroxidase (GPX), glutathione-Stransferase (GST), and catalase (CAT), and the non-enzymatic low molecular compounds like ascorbic acid (AA), reduced glutathione (GSH), α-tocopherol, carotenoids, phenolics, flavonoids, and proline (Gill and Tuteja, 2010; Miller et al., 2010; Gill et al., 2011)....

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Journal ArticleDOI
TL;DR: The description outlined here facilitates the understanding of factors that favour mitochondrial ROS production and develops better methods to measure mitochondrial O2•− and H2O2 formation in vivo, as uncertainty about these values hampers studies on the role of mitochondrial ROS in pathological oxidative damage and redox signalling.
Abstract: The production of ROS (reactive oxygen species) by mammalian mitochondria is important because it underlies oxidative damage in many pathologies and contributes to retrograde redox signalling from the organelle to the cytosol and nucleus. Superoxide (O2•−) is the proximal mitochondrial ROS, and in the present review I outline the principles that govern O2•− production within the matrix of mammalian mitochondria. The flux of O2•− is related to the concentration of potential electron donors, the local concentration of O2 and the second-order rate constants for the reactions between them. Two modes of operation by isolated mitochondria result in significant O2•− production, predominantly from complex I: (i) when the mitochondria are not making ATP and consequently have a high Δp (protonmotive force) and a reduced CoQ (coenzyme Q) pool; and (ii) when there is a high NADH/NAD+ ratio in the mitochondrial matrix. For mitochondria that are actively making ATP, and consequently have a lower Δp and NADH/NAD+ ratio, the extent of O2•− production is far lower. The generation of O2•− within the mitochondrial matrix depends critically on Δp, the NADH/NAD+ and CoQH2/CoQ ratios and the local O2 concentration, which are all highly variable and difficult to measure in vivo. Consequently, it is not possible to estimate O2•− generation by mitochondria in vivo from O2•−-production rates by isolated mitochondria, and such extrapolations in the literature are misleading. Even so, the description outlined here facilitates the understanding of factors that favour mitochondrial ROS production. There is a clear need to develop better methods to measure mitochondrial O2•− and H2O2 formation in vivo, as uncertainty about these values hampers studies on the role of mitochondrial ROS in pathological oxidative damage and redox signalling.

6,371 citations


"Reactive oxygen species (ROS) and r..." refers background in this paper

  • ...In Complex III, ubiquinone in its fully reduced form donates an electron to Cytochrome c1 leaving behind an unstable ubisemiquinone semi-radical which favors leakage of electrons to O2, thereby generating O •− 2 (Murphy, 2009)....

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Journal ArticleDOI
TL;DR: This review describes the main mitochondrial sources of reactive species and the antioxidant defences that evolved to prevent oxidative damage in all the mitochondrial compartments and discusses various physiological and pathological scenarios resulting from an increased steady state concentration of mitochondrial oxidants.
Abstract: The reduction of oxygen to water proceeds via one electron at a time. In the mitochondrial respiratory chain, Complex IV (cytochrome oxidase) retains all partially reduced intermediates until full reduction is achieved. Other redox centres in the electron transport chain, however, may leak electrons to oxygen, partially reducing this molecule to superoxide anion (O2−•). Even though O2−• is not a strong oxidant, it is a precursor of most other reactive oxygen species, and it also becomes involved in the propagation of oxidative chain reactions. Despite the presence of various antioxidant defences, the mitochondrion appears to be the main intracellular source of these oxidants. This review describes the main mitochondrial sources of reactive species and the antioxidant defences that evolved to prevent oxidative damage in all the mitochondrial compartments. We also discuss various physiological and pathological scenarios resulting from an increased steady state concentration of mitochondrial oxidants.

4,282 citations


"Reactive oxygen species (ROS) and r..." refers background in this paper

  • ...This reverse flow of electrons is controlled by ATP hydrolysis (Turrens, 2003)....

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