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

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

Over-production of reactive oxygen species (ROS) in plants under stress conditions is a common phenomenon. Plants tend to counter this problem through their ability to synthesize ROS neutralizing substances including non-enzymatic and enzymatic antioxidants. In this context, ascorbic acid (AsA) is one of the universal non-enzymatic antioxidants having substantial potential of not only scavenging ROS, but also modulating a number of fundamental functions in plants both under stress and non-stress conditions. In the present review, the role of AsA, its biosynthesis, and cross-talk with different hormones have been discussed comprehensively. Furthermore, the possible involvement of AsA-hormone crosstalk in the regulation of several key physiological and biochemical processes like seed germination, photosynthesis, floral induction, fruit expansion, ROS regulation and senescence has also been described. A simplified and schematic AsA biosynthetic pathway has been drawn, which reflects key intermediates involved therein. This could pave the way for future research to elucidate the modulation of plant AsA biosynthesis and subsequent responses to environmental stresses. Apart from discussing the role of different ascorbate peroxidase isoforms, the comparative role of two key enzymes, ascorbate peroxidase (APX) and ascorbate oxidase (AO) involved in AsA metabolism in plant cell apoplast is also discussed particularly focusing on oxidative stress perception and amplification. Limited progress has been made so far in terms of developing transgenics which could over-produce AsA. The prospects of generation of transgenics overexpressing AsA related genes and exogenous application of AsA have been discussed at length in the review.

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Ascorbic Acid-A Potential Oxidant Scavenger and Its Role in Plant Development and Abiotic Stress Tolerance.

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.

https://www.mdpi.com/1422-0067/21/22/8695/pdf

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

This study was carried out to assess the influence of trehalose, a non-reducing disaccharide involved in improving plant stress tolerance, on two cultivars (Hysun 33 and FH 598) of sunflower (Helianthus annuus L.) grown under control and drought stress conditions. At pre-flowering stage, varying concentrations (10, 20 and 30 mM) of trehalose were applied to the foliage. Drought stress significantly suppressed the plant growth, total soluble proteins, chlorophyll, achene yield per plant, oil percentage, organic contents, as well as oil palmitic and linoleic acids in both sunflower cultivars. External application of trehalose significantly reduced RMP (relative membrane permeability), and the accumulation of H2 O2 (hydrogen peroxide), while a considerable improvement was recorded in shoot fresh and shoot and root dry weights, total soluble proteins, glycinebetaine, AsA (ascorbic acid), total phenolics, achene yield per plant, oil contents, inorganic and organic contents, and the activities of catalase (CAT), superoxide dismutase (SOD) and peroxidase (POD) enzymes under water-limited regimes. The cultivar Hysun 33 was superior to the other cultivar in plant growth, RMP, glycinebetaine, proline, achene yield per plant, oil contents, and palmitic and linoleic acids. Overall, foliar-applied trehalose improved plant growth, oxidative defense system, yield and oil composition of sunflower under drought stress conditions.

Impact of exogenously applied trehalose on leaf biochemistry, achene yield and oil composition of sunflower under drought stress.

A study was conducted to find out the role of ascorbic acid (AsA) in modulating growth and different physio-biochemical attributes of canola plants under well-watered as well as water-deficit conditions. Drought stress imposed on 60 % field capacity significantly decreased the shoot and root fresh and dry weights, leaf chlorophyll contents, shoot and root P, root K+, and activity of CAT enzyme, while increased chlorophyll a/b contents, MDA, NPQ, leaf total phenolics, free proline and GB contents in both canola cultivars. Foliar-applied varying levels (50, 100 and 150 mg L−1) of AsA enhanced shoot and root fresh and root dry weights, qN, NPQ, shoot and root P, AsA as well as the activity of POD enzyme particularly under drought stress conditions. Of both canola cultivars, cv. Dunkeld was higher in shoot fresh weights, ETR and F
 v
 /F
 m, MDA, proline and GB contents, and POD activity, however, cv. Cyclone in total phenolics and qN under well-watered and water-deficit conditions. Overall, the foliar-applied AsA had a positive effect, though not marked, on salt sensitive cv. Cyclone in terms of improved growth and other attributes, whereas exogenously applied AsA had a non-significant effect on relatively salt tolerant cv. Dunkeld.

Synergistic effects of drought and ascorbic acid on growth, mineral nutrients and oxidative defense system in canola ( Brassica napus L.) plants

The effects of seed priming by ascorbic acid on some morphological and biochemical aspects of rapeseed (Brassica napus L.) under drought stress condition

The role of ascorbic acid as a key molecule antioxidant involved in biotic and abiotic stress has been well described. Thus, in order to study the effects of ascorbic acid on drought tolerance and some morphological and physiological characteristics of safflower (Carthamus tinctorius L.) under drought stress condition an experiment was conducted. The experimental design was factorial with two factors arranged in a completely randomized design (CRD), with three replications. The first factor was drought stress on 5 levels (control, -4, -6 , -8 and -12 bar) that was carried out by PEG 6000 and the second factor was ascorbic acid solution on 4 density (control, 55, 110 and 165 μm). The results showed that with increase in drought stress, germination percentage, shoot length, root length, seedling fresh weight, seedling dry weight and vigor index significantly decreased whereas catalase and peroxidase activity increased as compared to control with enhancement of drought stress. In general, priming with ascorbic acid significantly relived the harsh effects of drought stress on seedling growth, catalase and peroxidase activity of safflower and it seems that ascorbic acid was able to enhance the tolerant ability of the plant to drought stress.

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The effects of seed priming with ascorbic acid on drought tolerance and some morphological and physiological characteristics of safflower (Carthamus tinctorius L.).

Maryam Farzanian

Papers

The effects of seed priming by ascorbic acid on some morphological and biochemical aspects of rapeseed (Brassica napus L.) under drought stress condition

The effects of seed priming with ascorbic acid on drought tolerance and some morphological and physiological characteristics of safflower (Carthamus tinctorius L.).