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.

Arsenic uptake, toxicity, detoxification, and speciation in plants : physiological, biochemical, and molecular aspects

Climate change is an invisible, silent killer with calamitous effects on living organisms As the sessile organism, plants experience a diverse array of abiotic stresses during ontogenesis The relentless climatic changes amplify the intensity and duration of stresses, making plants dwindle to survive Plants convert 1-2% of consumed oxygen into reactive oxygen species (ROS), in particular, singlet oxygen (1O2), superoxide radical (O2•-), hydrogen peroxide (H2O2), hydroxyl radical (•OH), etc as a byproduct of aerobic metabolism in different cell organelles such as chloroplast, mitochondria, etc The regulatory network comprising enzymatic and non-enzymatic antioxidant systems tends to keep the magnitude of ROS within plant cells to a non-damaging level However, under stress conditions, the production rate of ROS increases exponentially, exceeding the potential of antioxidant scavengers instigating oxidative burst, which affects biomolecules and disturbs cellular redox homeostasis ROS are similar to a double-edged sword; and, when present below the threshold level, mediate redox signaling pathways that actuate plant growth, development, and acclimatization against stresses The production of ROS in plant cells displays both detrimental and beneficial effects However, exact pathways of ROS mediated stress alleviation are yet to be fully elucidated Therefore, the review deposits information about the status of known sites of production, signaling mechanisms/pathways, effects, and management of ROS within plant cells under stress In addition, the role played by advancement in modern techniques such as molecular priming, systems biology, phenomics, and crop modeling in preventing oxidative stress, as well as diverting ROS into signaling pathways has been canvassed

Abiotic Stress and Reactive Oxygen Species: Generation, Signaling, and Defense Mechanisms

Nitric oxide alleviates silver nanoparticles (AgNps)-induced phytotoxicity in Pisum sativum seedlings.

Phenolic composition and antioxidant potential of grain legume seeds: A review.

Nitric oxide regulates plant responses to drought, salinity, and heavy metal stress

The adverse effects of arsenic (As) toxicity on seedling growth, root and shoot anatomy, chlorophyll and carotenoid contents, root oxidizability (RO), antioxidant enzyme activities, H2O2 content, lipid peroxidation and electrolyte leakage (EL%) in common bean (Phaseolus vulgaris L.) were investigated. The role of exogenous nitric oxide (NO) in amelioration of As-induced inhibitory effect was also evaluated using sodium nitroprusside (100 μM SNP) as NO donor and 2-(4-carboxy-2-phenyl)-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide (200 μM PTIO) as NO scavenger in different combinations with 50 μM As. As-induced growth inhibition was associated with marked anomalies in anatomical features, reduction in pigment composition, increased RO and severe perturbations in antioxidant enzyme activities. While activity of superoxide dismutase and catalase increased, levels of ascorbate peroxidase, dehydroascorbate reductase and glutathione reductase decreased significantly and guaiacol peroxidase remained normal. The over-accumulation of H2O2 content along with high level of lipid peroxidation and electrolyte leakage indicates As-induced oxidative damage in P. vulgaris seedlings with more pronounced effect on the roots than the shoots. Exogenous addition of NO significantly reversed the As-induced oxidative stress, maintaining H2O2 in a certain level through balanced alterations of antioxidant enzyme activities. The role of NO in the process of amelioration has ultimately been manifested by significant reduction of membrane damage and improvement of growth performance in plants grown on As + SNP media. Onset of oxidative stress was more severe after addition of PTIO, which confirms the protective role of NO against As-induced oxidative damage in P. vulgaris seedlings.

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Arsenic-induced oxidative stress in the common bean legume, Phaseolus vulgaris L. seedlings and its amelioration by exogenous nitric oxide

The effect of calcium (Ca) on lentil (Lens culinaris Medic.) seedlings exposed to cadmium (Cd) stress was studied by investigating plant growth and antioxidant enzyme activities. Plants were grown for 14 days in full-strength Hoagland nutrient media supplemented with Cd concentrations of 0, 10, 20, and 40 μM, and on corresponding medium supplied with 5 mM Ca(NO3)2 prior to Cd addition. Increasing Cd led to accumulation of metal and reduced the fresh weight of the shoots more strongly than that of the roots. Cd concentrations of 20 and 40 μM were selected to study its toxic effect on seedlings. Activities of superoxide dismutase, ascorbate peroxidase, catalase, dehydroascorbate reductase, and glutathione reductase decreased at much higher magnitude in the shoots than those observed in the roots under Cd exposure. Failure of antioxidant defense in scavenging of reactive oxygen species was evidenced by abnormal rise in H2O2, resulting in enhancement of lipid peroxidation and membrane electrolyte leakage as the marks of Cd-induced oxidative stress in lentil seedlings. Ca priming in the media significantly reduced the Cd accumulation and considerably alleviated the adverse impact of Cd treatment by modulating the antioxidant enzyme activity. Mitigation of Cd-induced stress by Ca application was strongly suggested by declining levels of H2O2 and consequent lowering of oxidative damage of membrane. Consequently, this enhanced fresh mass of plant parts as the sign of Ca-mediated normal growth in Cd-treated lentil seedlings.

Exogenous calcium alleviates the impact of cadmium-induced oxidative stress in Lens culinaris medic. Seedlings through modulation of antioxidant enzyme activities

The effects of arsenic (As) on growth and antioxidant metabolism of fenugreek (Trigonella foenum-graecum L. cv. Azad) plants were studied using 10, 20, and 30 mg As/kg of soil in a pot experiment under controlled conditions. The length and dry weights of shoots and roots, photosynthetic traits, and grain yield components exhibited a significant increase over control (0 mg As/kg) at As20 but decreased markedly at As30. The cause of this completely reverse response of plant growth between As20 and As30 was investigated in the backdrop of H2O2 metabolism by analyzing responses of three prominent antioxidant enzymes, namely superoxide dismutase (SOD), ascorbate peroxidase (APX), and catalase (CAT) along with cellular ascorbate pool and its redox state. Despite a significant increase in the H2O2 content in both As20 and As30 plants, the former, unlike As30 plants, did not experience any type of As-induced oxidative stress (membrane lipid peroxidation, electrolyte leakage). Normal to high levels of leaf APX, CAT, and redox pool of ascorbate effectively balanced the elevated SOD activity at As20, maintaining the H2O2 concentration higher than control but obviously in favor of As20 plant growth. By contrast, soil amendment with phosphorus (200 mg P/kg) at As30 prevented As-induced oxidative stress through the reduction of the H2O2 level even below As0. The increased enzyme activity was mainly due to the induction of unique Cu/Zn, Fe, and Mn isoforms of SOD and APX-3/APX-4 and/or their increased expression in coordination with CAT. The detection of novel isoforms suggests a strong response of H2O2-metabolizing enzymes against As-induced oxidative stress in fenugreek.

Arsenic-induced changes in growth and antioxidant metabolism of fenugreek

Response of sulfate transporters, thiol metabolism, and antioxidant defense system was studied in roots of two lentil (Lens culinaris Medik.) genotypes grown in arsenic (10, 25, and 40 μM As(V))-supplemented nutrient solution, and significant changes compared to control (0 μM As(V)) were observed mainly at 25 and 40 μM. In L 414, high glutathione (GSH) redox (0.8-0.9) was maintained with elevated thiol synthesis, powered by transcriptional up-regulation of LcSultr1;1 and LcSultr1;2 sulfate transporters and significant induction of LcSAT1;1 and LcSAT1;2 (serine acetyltransferase), OAS-TL (O-acetylserine(thiol)-lyase), γ-ECS (γ-glutamylcysteine synthetase), and PCS (phytochelatin synthase) genes predominantly within 12-24 h of As exposure at 25 μM and within 6-12 h at 40 μM. This thiolic potency in L 414 roots was effectively complemented by up-regulation of gene expressions and consequent enhanced activities of superoxide dismutase, ascorbate peroxidase (APX), dehydroascorbate reductase, glutathione reductase (GR), and glutathione-S-transferase (GST) isoforms at 25 and 40 μMAs, efficiently scavenging excess reactive oxygen species to prevent onset of As-induced oxidative stress and consequent inhibition of root growth in L 414. In contrast, down-regulation of vital sulfate-uptake transporters as well as entire thiol-metabolizing system and considerably low APX, GST, and GR expressions in DPL 59 not only resulted in reduced GSH redox but also led to over-accumulation of H2O2. This triggered membrane lipid peroxidations as the marks of As-induced oxidative damage. Results indicated coordinated response of thiol-metabolism and antioxidant defense in conferring As-tolerance in lentil, and GSH is the key point in this cascade.

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Retraction Note to: Coordinated response of sulfate transport, cysteine biosynthesis, and glutathione-mediated antioxidant defense in lentil (Lens culinaris Medik.) genotypes exposed to arsenic.

Background: Both lentil (Lens culinaris Medik.) and grass pea (Lathyrus sativus L.) in the family Fabaceae are two important cool-season food legumes, often experiencing water stress conditions during growth and maturity. Objective: The present study was undertaken to ascertain the response of these two crops under different water stress regimes. Materials and Methods: Different morpho-physiological and biochemical parameters were studied in a pot experiment under controlled environmental conditions. Along with control (proper irrigation, 0 stress), three sets of plants were subjected to mild (6 d), moderate (13 d) and severe (20 d) water stress by withholding irrigation at the appropriate time. Results: Compared with control, plant growth traits and seed yield components reduced significantly in both crops with increasing period of water stress, resulting in lowering of dry mass with more severe effect on lentil compared with grass pea. Foliar Relative Water Content (RWC) (%), K + /Na + ratio, chlorophyll (chl) a, chl a/b ratio, stomatal conductance and net photosynthetic rate declined considerably in both crops under water stress. Leaf-free proline level increased significantly in both crops, but it decreased markedly in nodules of lentil and remained unchanged in grass pea. Nodulation was also affected due to water stress. The impairment in growth traits and physio-biochemical parameters under water stress was manifested in reduction of drought tolerance efficiency of both crops. Conclusion: Impact of water stress was more severe on lentil compared with grass pea, and modulation of growth traits signified necessity of a detailed strategy in breeding of food legumes under water stress.

Dibyendu Talukdar

Papers

Arsenic-induced oxidative stress in the common bean legume, Phaseolus vulgaris L. seedlings and its amelioration by exogenous nitric oxide

Exogenous calcium alleviates the impact of cadmium-induced oxidative stress in Lens culinaris medic. Seedlings through modulation of antioxidant enzyme activities

Arsenic-induced changes in growth and antioxidant metabolism of fenugreek

Retraction Note to: Coordinated response of sulfate transport, cysteine biosynthesis, and glutathione-mediated antioxidant defense in lentil (Lens culinaris Medik.) genotypes exposed to arsenic.

Comparative morpho-physiological and biochemical responses of lentil and grass pea genotypes under water stress