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      Are Flavonoids Effective Antioxidants in Plants? Twenty Years of Our Investigation

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          Abstract

          Whether flavonoids play significant antioxidant roles in plants challenged by photooxidative stress of different origin has been largely debated over the last few decades. A critical review of the pertinent literature and our experimentation as well, based on a free-of-scale approach, support an important antioxidant function served by flavonoids in plants exposed to a wide range of environmental stressors, the significance of which increases with the severity of stress. On the other side, some questions need conclusive answers when the putative antioxidant functions of plant flavonoids are examined at the level of both the whole-cell and cellular organelles. This partly depends upon a conclusive, robust, and unbiased definition of “a plant antioxidant”, which is still missing, and the need of considering the subcellular re-organization that occurs in plant cells in response to severe stress conditions. This likely makes our deterministic-based approach unsuitable to unveil the relevance of flavonoids as antioxidants in extremely complex biological systems, such as a plant cell exposed to an ever-changing stressful environment. This still poses open questions about how to measure the occurred antioxidant action of flavonoids. Our reasoning also evidences the need of contemporarily evaluating the changes in key primary and secondary components of the antioxidant defense network imposed by stress events of increasing severity to properly estimate the relevance of the antioxidant functions of flavonoids in an in planta situation. In turn, this calls for an in-depth analysis of the sub-cellular distribution of primary and secondary antioxidants to solve this still intricate matter.

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          ROS Are Good.

          Reactive oxygen species (ROS) are thought to play a dual role in plant biology. They are required for many important signaling reactions, but are also toxic byproducts of aerobic metabolism. Recent studies revealed that ROS are necessary for the progression of several basic biological processes including cellular proliferation and differentiation. Moreover, cell death-that was previously thought to be the outcome of ROS directly killing cells by oxidation, in other words via oxidative stress-is now considered to be the result of ROS triggering a physiological or programmed pathway for cell death. This Opinion focuses on the possibility that ROS are beneficial to plants, supporting cellular proliferation, physiological function, and viability, and that maintaining a basal level of ROS in cells is essential for life.
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            Reactive oxygen species, abiotic stress and stress combination.

            Reactive oxygen species (ROS) play a key role in the acclimation process of plants to abiotic stress. They primarily function as signal transduction molecules that regulate different pathways during plant acclimation to stress, but are also toxic byproducts of stress metabolism. Because each subcellular compartment in plants contains its own set of ROS-producing and ROS-scavenging pathways, the steady-state level of ROS, as well as the redox state of each compartment, is different at any given time giving rise to a distinct signature of ROS levels at the different compartments of the cell. Here we review recent studies on the role of ROS in abiotic stress in plants, and propose that different abiotic stresses, such as drought, heat, salinity and high light, result in different ROS signatures that determine the specificity of the acclimation response and help tailor it to the exact stress the plant encounters. We further address the role of ROS in the acclimation of plants to stress combination as well as the role of ROS in mediating rapid systemic signaling during abiotic stress. We conclude that as long as cells maintain high enough energy reserves to detoxify ROS, ROS is beneficial to plants during abiotic stress enabling them to adjust their metabolism and mount a proper acclimation response.
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              Flavonoids as antioxidants in plants: location and functional significance.

              Stress-responsive dihydroxy B-ring-substituted flavonoids have great potential to inhibit the generation of reactive oxygen species (ROS) and reduce the levels of ROS once they are formed, i.e., to perform antioxidant functions. These flavonoids are located within or in the proximity of centers of ROS generation in severely stressed plants. Efficient mechanisms have been recently identified for the transport of flavonoids from the endoplasmic reticulum, the site of their biosynthesis, to different cellular compartments. The mechanism underlying flavonoid-mediated ROS reduction in plants is still unclear. 'Antioxidant' flavonoids are found in the chloroplast, which suggests a role as scavengers of singlet oxygen and stabilizers of the chloroplast outer envelope membrane. Dihydroxy B-ring substituted flavonoids are present in the nucleus of mesophyll cells and may inhibit ROS-generation making complexes with Fe and Cu ions. The genes that govern the biosynthesis of antioxidant flavonoids are present in liverworts and mosses and are mostly up-regulated as a consequence of severe stress. This suggests that the antioxidant flavonoid metabolism is a robust trait of terrestrial plants. Vacuolar dihydroxy B-ring flavonoids have been reported to serve as co-substrates for vacuolar peroxidases to reduce H(2)O(2) escape from the chloroplast, following the depletion of ascorbate peroxidase activity. Antioxidant flavonoids may effectively control key steps of cell growth and differentiation, thus acting regulating the development of the whole plant and individual organs. Copyright © 2012 Elsevier Ireland Ltd. All rights reserved.
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                Author and article information

                Journal
                Antioxidants (Basel)
                Antioxidants (Basel)
                antioxidants
                Antioxidants
                MDPI
                2076-3921
                09 November 2020
                November 2020
                : 9
                : 11
                : 1098
                Affiliations
                [1 ]Institute of Applied Physics ‘Carrara’, National Research Council of Italy (CNR), Via Madonna del Piano 10, Sesto F.no, I-50019 Florence, Italy; g.agati@ 123456ifac.cnr.it
                [2 ]Institute for Sustainable Plant Protection, National Research Council of Italy (CNR), Via Madonna del Piano 10, I-50019, Sesto F.no, Florence, Italy; cecilia.brunetti@ 123456ipsp.cnr.it (C.B.); federico.sebastiani@ 123456ipsp.cnr.it (F.S.)
                [3 ]Department of Agriculural and Environmental Sciences - Production, Landscape, Agroenergy, University of Milan, Via Celoria 2, I-20133 Milan, Italy; alessio.fini@ 123456unimi.it
                [4 ]Department of Agriculture, Food, Environment and Forestry, University of Florence, Viale delle Idee 30, Sesto F.no, I-50019 Florence, Italy; antonella.gori@ 123456unifi.it
                [5 ]Department of Agriculture, Food and Environment, University of Pisa, Via del Borghetto 80, I-56124 Pisa, Italy; lucia.guidi@ 123456unipi.it (L.G.); marco.landi@ 123456unipi.it (M.L.)
                Author notes
                [* ]Correspondence: massimiliano.tattini@ 123456ipsp.cnr.it ; Tel.: +39-055-644-202
                Author information
                https://orcid.org/0000-0003-0855-9389
                https://orcid.org/0000-0002-8531-6076
                https://orcid.org/0000-0002-7304-7526
                https://orcid.org/0000-0002-2472-720X
                https://orcid.org/0000-0003-0121-0715
                https://orcid.org/0000-0001-5434-8860
                Article
                antioxidants-09-01098
                10.3390/antiox9111098
                7695271
                33182252
                75539f9c-925d-4893-903a-5e473ccfdfca
                © 2020 by the authors.

                Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license ( http://creativecommons.org/licenses/by/4.0/).

                History
                : 12 October 2020
                : 06 November 2020
                Categories
                Review

                antioxidant enzymes,cytoplasm-located flavonoids,early land plants,flavonols,hydrogen peroxide,photoprotection,reactive oxygen species,uv-b radiation,vacuolar flavonoids

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