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The stationary life of plants has led to the evolution of a complex gridded antioxidant defence system constituting numerous enzymatic components, playing a crucial role in overcoming various stress conditions. Mainly, these plant enzymes are superoxide dismutase (SOD), catalase (CAT), peroxidase (POX), glutathione peroxidase (GPX), glutathione reductase (GR), glutathione S-transferases (GST), ascorbate peroxidase (APX), monodehydroascorbate reductase (MDHAR), and dehydroascorbate reductase (DHAR), which work as part of the antioxidant defence system. These enzymes together form a complex set of mechanisms to minimise, buffer, and scavenge the reactive oxygen species (ROS) efficiently. The present review is aimed at articulating the current understanding of each of these enzymatic components, with special attention on the role of each enzyme in response to the various environmental, especially abiotic stresses, their molecular characterisation, and reaction mechanisms. The role of the enzymatic defence system for plant health and development, their significance, and cross-talk mechanisms are discussed in detail. Additionally, the application of antioxidant enzymes in developing stress-tolerant transgenic plants are also discussed.

Plants are exposed to various environmental stresses in their lifespan that threaten their survival. Reactive oxygen species (ROS), the byproducts of aerobic metabolism, are essential signalling molecules in regulating multiple plant developmental processes as well as in reinforcing plant tolerance to biotic and abiotic stimuli. However, intensified environmental challenges such as salinity, drought, UV irradiation, and heavy metals usually interfere with natural ROS metabolism and homeostasis, thus aggravating ROS generation excessively and ultimately resulting in oxidative stress. Cellular damage is confined to the degradation of biomolecular structures, including carbohydrates, proteins, lipids, pigments, and DNA. The nature of the double-edged function of ROS as a secondary messenger or harmful oxidant has been attributed to the degree of existing balance between cellular ROS production and ROS removal machinery. The activities of enzyme-based antioxidants, catalase (CAT, EC 1.11.1.6), monodehydroascorbate reductase (MDHAR, E.C.1.6.5.4), dehydroascorbate reductase (DHAR, EC 1.8.5.1), superoxide dismutase (SOD, EC 1.15.1.1), ascorbate peroxidase (APX, EC 1.11.1.11), glutathione reductase (GR, EC 1.6.4.2), and guaiacol peroxidase (GPX, EC 1.11.1.7); and non-enzyme based antioxidant molecules, ascorbate (AA), glutathione (GSH), carotenoids, α-tocopherol, prolines, flavonoids, and phenolics, are indeed parts of the defensive strategies developed by plants to scavenge excess ROS and to maintain cellular redox homeostasis during oxidative stress. This review briefly summarises current knowledge on enzymatic and non-enzymatic antioxidant machinery in plants. Moreover, additional information about the beneficial impact of the microbiome on countering abiotic/biotic stresses in association with roots and plant tissues has also been provided.

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.

Tomato is an important vegetable that is highly sensitive to drought (DR) stress which impairs the development of tomato seedlings. Recently, melatonin (ME) has emerged as a nontoxic, regulatory biomolecule that regulates plant growth and enhances the DR tolerance mechanism in plants. The present study was conducted to examine the defensive role of ME in photosynthesis, root architecture, and the antioxidant enzymes’ activities of tomato seedlings subjected to DR stress. Our results indicated that DR stress strongly suppressed growth and biomass production, inhibited photosynthesis, negatively affected root morphology, and reduced photosynthetic pigments in tomato seedlings. Per contra, soluble sugars, proline, and ROS (reactive oxygen species) were suggested to be improved in seedlings under DR stress. Conversely, ME (100 µM) pretreatment improved the detrimental-effect of DR by restoring chlorophyll content, root architecture, gas exchange parameters and plant growth attributes compared with DR-group only. Moreover, ME supplementation also mitigated the antioxidant enzymes [APX (ascorbate peroxidase), CAT (catalase), DHAR (dehydroascorbate reductase), GST (glutathione S-transferase), GR (glutathione reductase), MDHAR (monodehydroascorbate reductase), POD (peroxidase), and SOD (superoxide dismutase)], non-enzymatic antioxidant [AsA (ascorbate), DHA (dehydroascorbic acid), GSH (glutathione), and GSSG, (oxidized glutathione)] activities, reduced oxidative damage [EL (electrolyte leakage), H2O2 (hydrogen peroxide), MDA (malondialdehyde), and O2•− (superoxide ion)] and osmoregulation (soluble sugars and proline) of tomato seedlings, by regulating gene expression for SOD, CAT, APX, GR, POD, GST, DHAR, and MDHAR. These findings determine that ME pretreatment could efficiently improve the seedlings growth, root characteristics, leaf photosynthesis and antioxidant machinery under DR stress and thereby increasing the seedlings’ adaptability to DR stress.

Experiments were carried out to investigate the role of nitric oxide (NO) in ameliorating the negative effects of cadmium stress in tomato seedlings. Plants treated with cadmium (CdCl2, 150 μM) showed reduced growth, biomass yield, pigment content, chlorophyll fluorescence, and gas exchange parameters. Exogenous application of NO donor (sodium nitroprusside) with nutrient solution protected chlorophyll pigments, restored chlorophyll fluorescence and gas exchange parameters, and caused significant enhancements in growth and biomass yield. Cadmium triggered the synthesis of proline and glycine betaine; however, application of NO caused further enhancement of their accumulation, reflecting an obvious amelioration of the cadmium-induced decline in relative water content. Activities of the antioxidant enzymes superoxide dismutase, catalase, ascorbate peroxidase, and glutathione reductase, monodehydroascorbate reductase, dehydroascorbate reductase, and other enzymatic activities of ascorbate-glutathione cycle were enhanced following the application of NO, as compared with those in untreated seedlings under control and cadmium stress conditions. NO increased the flavonoid and total phenol content in Cd-stressed tomato plants. Moreover, NO application restricted the uptake of cadmium and enhanced the accumulation of nutrients in different parts of tomato plants. On the basis of the findings of the present study, we propose that NO has a potential role as a growth promoter for tomato under cadmium stress.

Background Salinity-alkalinity stress is one of the major abiotic stresses affecting plant growth and development. γ-Aminobutyrate (GABA) is a non-protein amino acid that functions in stress tolerance. However, the interactions between cellular redox signaling and chlorophyll (Chl) metabolism involved in GABA-induced salinity-alkalinity stress tolerance in plants remains largely unknown. Here, we investigated the role of GABA in perceiving and regulating chlorophyll biosynthesis and oxidative stress induced by salinity-alkalinity stress in muskmelon leaves. We also evaluated the effects of hydrogen peroxide (H 2 O 2 ), glutathione (GSH), and ascorbate (AsA) on GABA-induced salinity-alkalinity stress tolerance. Results Salinity-alkalinity stress increased malondialdehyde (MDA) content, relative electrical conductivity (REC), and the activities of superoxide dismutase (SOD), ascorbate peroxidase (APX) and dehydroascorbate reductase (DHAR). Salinity-alkalinity stress decreased shoot dry and fresh weight and leaf area, reduced glutathione and ascorbate (GSH and AsA) contents, activities of glutathione reductase (GR) and monodehydroascorbate reductase (MDAR). By contrast, pretreatment with GABA, H 2 O 2 , GSH, or AsA significantly inhibited these salinity-alkalinity stress-induced effects. The ability of GABA to relieve salinity-alkalinity stress was significantly reduced when the production of endogenous H 2 O 2 was inhibited, but was not affected by inhibiting endogenous AsA and GSH production. Exogenous GABA induced respiratory burst oxidase homologue D ( RBOHD ) genes expression and H 2 O 2 accumulation under normal conditions but reduced the H 2 O 2 content under salinity-alkalinity stress. Salinity-alkalinity stress increased the accumulation of the chlorophyll synthesis precursors glutamate (Glu), δ-aminolevulinic acid (ALA), porphobilinogen (PBG), uroporphyrinogen III (URO III), Mg-protoporphyrin IX (Mg-proto IX), protoporphyrin IX (Proto IX), protochlorophyll (Pchl), thereby increasing the Chl content. Under salinity-alkalinity stress, exogenous GABA increased ALA content, but reduced the contents of Glu, PBG, URO III, Mg-proto IX, Proto IX, Pchl, and Chl. However, salinity-alkalinity stress or GABA treated plant genes expression involved in Chl synthesis had no consistent trends with Chl precursor contents. Conclusions Exogenous GABA elevated H 2 O 2 may act as a signal molecule, while AsA and GSH function as antioxidants, in GABA-induced salinity-alkalinity tolerance. These factors maintain membrane integrity which was essential for the ordered chlorophyll biosynthesis. Pretreatment with exogenous GABA mitigated salinity-alkalinity stress caused excessive accumulation of Chl and its precursors, to avoid photooxidation injury.

Melatonin is a multifunctional signaling molecule that is ubiquitously distributed in different parts of a plant and responsible for stimulating several physio-chemical responses to adverse environmental conditions. In this review, we show that, although plants are able to biosynthesize melatonin, the exogenous application of melatonin to various crops can improve plant growth and development in response to various abiotic and biotic stresses (e.g., drought, unfavorable temperatures, high salinity, heavy metal contamination, acid rain, and combined stresses) by regulating antioxidant machinery of plants. Current knowledge suggests that exogenously applied melatonin can enhance the stress tolerance of plants by regulating both the enzymatic and non-enzymatic antioxidant defense systems. Enzymic antioxidants upregulated by exogenous melatonin include superoxide dismutase, catalase, glutathione peroxidase, and enzymes involved in the ascorbate–glutathione cycle (ascorbate peroxidase, monodehydroascorbate reductase, dehydroascorbate reductase, and glutathione reductase), whereas levels of non-enzymatic antioxidants such as ascorbate, reduced glutathione, carotenoids, tocopherols, and phenolics are also higher under stress conditions. The enhanced antioxidant system consequently exhibits lower lipid peroxidation and greater plasma membrane integrity when under stress. However, these responses vary greatly from crop to crop and depend on the intensity and type of stress, and most studies to date have been conducted under controlled conditions. This means that a wider range of crop field trials and detailed transcriptomic analysis are required to reveal the gene regulatory networks involved in the between melatonin, antioxidants, and abiotic stress.

With the rapid development of nanotechnology, the potential releases of nanoparticles (NPs) have drawn considerable attention. Plants are essential fundamental components of all ecosystems, and the interaction between NPs and plants is an indispensable aspect of the risk assessment. Originally, this review focuses on NP phytotoxicity, which is an important precondition to promote the application of nanotechnology and to avoid the potential ecological risks. Both enhancive and inhibitive effects of various NPs on different plants’ growth have been documented. In this paper, the influence factors of nanotoxicity and the mechanisms of these toxic effects are also summarized. Subsequently, the defense mechanisms are presented as well. Eventually, this review puts forward the prospects of research direction of the environmental behavior and the biological toxicity of NPs, hoping to bring new ideas to the further research on NP phytotoxicity.Abbreviations: 1O2: singlet oxygen; AA: ascorbate; ABA: abscisic acid; APX: ascorbate peroxidase; B-Ag NPs: biosynthesized silver nanoparticles; Bt: Bacillus thuringiensis; CAT: catalase; CDs: carbon dots; Cu/Zn-SOD: copper/Zinc SOD; DHA: dehydroascorbate; DHAR: dehydroascorbate reductase; GA: gibberellic acid: GA-Ag NPs: coated silver nanoparticles; GPOX: guaiacol peroxidase; GPX: glutathione peroxidase; GR: glutathione reductase; GSH: glutathione; GSSG: oxidized glutathione; GST: glutathione S-transferases; •OH: hydroxyl radical; HA: high amylose; IAA: indole-3-acetic acid; LA: low amylose; MA: medium amylose; MDA: malondialdehyde; MDHAR: monodehydroascorbate reductase; NPs: nanoparticles; : superoxide radical; POD: promoting peroxidase; ROS: reactive oxygen species; SOD: superoxide dismutase; t-ZR: trans-zeatin-riboside

Background Microbes isolated from hyperaccumulating plants have been reported to be effective in achieving higher phytoextraction efficiency. The plant growth-promoting bacteria (PGPB) SaMR12 from the cadmium (Cd)/zinc hyperaccumulator Sedum alfredii Hance could promote the growth of a non-host plant, oilseed rape, under Cd stress. However, the effect of SaMR12 on Brasscia juncea antioxidative response under Cd exposure was still unclear. Results A hydroponic experiment was conducted to study the effects of Sphingomonas SaMR12 on its non-host plant Brassica juncea (L.) Czern. under four different Cd treatments. The results showed that SaMR12 could colonize and aggregate in the roots and then move to the shoots. SaMR12 inoculation promoted plant growth by up to 71% in aboveground biomass and 81% in root biomass over that of the non-inoculated plants. SaMR12-inoculated plants significantly enhanced root Cd accumulation in the 10 and 20 μM Cd treatments, with 1.72- and 0.86-fold increases, respectively, over that of the non-inoculated plants. SaMR12 inoculation not only decreased shoot hydrogen peroxide (H 2 O 2 ) content by up to 38% and malondialdehyde (MDA) content by up to 60% but also reduced proline content by 7–30% in shoots and 17–32% in roots compared to the levels in non-inoculated plants. Additionally, SaMR12 inoculation promoted the activities of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), and ascorbate peroxidase (APX) and facilitated the relative gene expression levels of dehydroascorbate reductase ( DHAR ) and glutathione reductase ( GR ) involved in the glutathione (GSH)-ascorbic acid (AsA) cycle. Conclusions The results demonstrated that, under Cd stress, SaMR12 inoculation could activate the antioxidative response of B. juncea by decreasing the concentrations of H 2 O 2 , MDA and proline, increasing the activities of antioxidative enzymes, and regulating the GSH-AsA cycle. These results provide a theoretical foundation for the potential application of hyperaccumulator endophytic bacteria as remediating agents to improve heavy metal tolerance within non-host plant species, which could further improve phytoextraction efficiency. Graphical abstract

The beneficial roles of melatonin (Mel) and gibberellic acid (GA3) in the biosynthesis of photosynthetic pigments, osmoregulation, and methylglyoxal (MG) detoxification and antioxidant system were studied in tomato (Solanum lycopersicum L. cv. Five Star) seedlings under NaCl stress. The exogenous application of Mel (100 µM) and GA3 (1.4 µM) together more efficiently affected growth performance of seedlings under salt stress. The decreased chlorophyll (Chl) degradation and Chl-degrading enzyme (chlorophyllase) activity in seedlings receiving Mel plus GA3 resulted in increased Chl content by upregulating Chl synthesizing enzyme (δ-aminolevulinic acid dehydratase) under salinity. Exogenous Mel plus GA3 suppressed the overproduction of reactive oxygen species (ROS; superoxide and hydrogen peroxide) and activity of glycolate oxidase. Application of Mel with GA3 reduced MG content by enhancing the activity of enzymes (glyoxalase I and glyoxalase II) involved in the MG detoxification system. Both Mel and GA3 together protected seedlings from ROS induced damage by regulating Δ1-pyrroline-5-carboxylate synthetase activity, and content of proline (Pro) and glycine betaine (GB). Seedlings receiving Mel + GA3 exhibited a substantial upregulated activity of catalase, ascorbate peroxidase, glutathione reductase, dehydroascorbate reductase, monodehydroascorbate reductase, glutathione peroxidase, polyphenol oxidase and lipoxygenase, and redox homeostasis that reduced oxidative damage induced by salinity. These outcomes advocate that Mel and GA3 played beneficial roles in Chl, Pro and GB biosynthesis, and improved redox homeostasis, and MG detoxification and antioxidant system under salt stress.