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      The Multiple Roles of Ascorbate in the Abiotic Stress Response of Plants: Antioxidant, Cofactor, and Regulator

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          Abstract

          Ascorbate (ASC) plays a critical role in plant stress response. The antioxidant role of ASC has been well-studied, but there are still several confusing questions about the function of ASC in plant abiotic stress response. ASC can scavenge reactive oxygen species (ROS) and should be helpful for plant stress tolerance. But in some cases, increasing ASC content impairs plant abiotic stress tolerance, whereas, inhibiting ASC synthesis or regeneration enhances plant stress tolerance. This confusing phenomenon indicates that ASC may have multiple roles in plant abiotic stress response not just as an antioxidant, though many studies more or less ignored other roles of ASC in plant. In fact, ACS also can act as the cofactor of some enzymes, which are involved in the synthesis, metabolism, and modification of a variety of substances, which has important effects on plant stress response. In addition, ASC can monitor and effectively regulate cell redox status. Therefore, we believe that ASC has atleast triple roles in plant abiotic stress response: as the antioxidant to scavenge accumulated ROS, as the cofactor to involve in plant metabolism, or as the regulator to coordinate the actions of various signal pathways under abiotic stress. The role of ASC in plant abiotic stress response is important and complex. The detail role of ASC in plant abiotic stress response should be analyzed according to specific physiological process in specific organ. In this review, we discuss the versatile roles of ASC in the response of plants to abiotic stresses.

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          Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants.

          Various abiotic stresses lead to the overproduction of reactive oxygen species (ROS) in plants which are highly reactive and toxic and cause damage to proteins, lipids, carbohydrates and DNA which ultimately results in oxidative stress. The ROS comprises both free radical (O(2)(-), superoxide radicals; OH, hydroxyl radical; HO(2), perhydroxy radical and RO, alkoxy radicals) and non-radical (molecular) forms (H(2)O(2), hydrogen peroxide and (1)O(2), singlet oxygen). In chloroplasts, photosystem I and II (PSI and PSII) are the major sites for the production of (1)O(2) and O(2)(-). In mitochondria, complex I, ubiquinone and complex III of electron transport chain (ETC) are the major sites for the generation of O(2)(-). The antioxidant defense machinery protects plants against oxidative stress damages. Plants possess very efficient enzymatic (superoxide dismutase, SOD; catalase, CAT; ascorbate peroxidase, APX; glutathione reductase, GR; monodehydroascorbate reductase, MDHAR; dehydroascorbate reductase, DHAR; glutathione peroxidase, GPX; guaicol peroxidase, GOPX and glutathione-S- transferase, GST) and non-enzymatic (ascorbic acid, ASH; glutathione, GSH; phenolic compounds, alkaloids, non-protein amino acids and α-tocopherols) antioxidant defense systems which work in concert to control the cascades of uncontrolled oxidation and protect plant cells from oxidative damage by scavenging of ROS. ROS also influence the expression of a number of genes and therefore control the many processes like growth, cell cycle, programmed cell death (PCD), abiotic stress responses, pathogen defense, systemic signaling and development. In this review, we describe the biochemistry of ROS and their production sites, and ROS scavenging antioxidant defense machinery. Copyright © 2010 Elsevier Masson SAS. All rights reserved.
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            ROS function in redox signaling and oxidative stress.

            Oxidative stress refers to elevated intracellular levels of reactive oxygen species (ROS) that cause damage to lipids, proteins and DNA. Oxidative stress has been linked to a myriad of pathologies. However, elevated ROS also act as signaling molecules in the maintenance of physiological functions--a process termed redox biology. In this review we discuss the two faces of ROS--redox biology and oxidative stress--and their contribution to both physiological and pathological conditions. Redox biology involves a small increase in ROS levels that activates signaling pathways to initiate biological processes, while oxidative stress denotes high levels of ROS that result in damage to DNA, protein or lipids. Thus, the response to ROS displays hormesis, given that the opposite effect is observed at low levels compared with that seen at high levels. Here, we argue that redox biology, rather than oxidative stress, underlies physiological and pathological conditions. Copyright © 2014 Elsevier Ltd. All rights reserved.
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              Plant hormone-mediated regulation of stress responses

              Background Being sessile organisms, plants are often exposed to a wide array of abiotic and biotic stresses. Abiotic stress conditions include drought, heat, cold and salinity, whereas biotic stress arises mainly from bacteria, fungi, viruses, nematodes and insects. To adapt to such adverse situations, plants have evolved well-developed mechanisms that help to perceive the stress signal and enable optimal growth response. Phytohormones play critical roles in helping the plants to adapt to adverse environmental conditions. The elaborate hormone signaling networks and their ability to crosstalk make them ideal candidates for mediating defense responses. Results Recent research findings have helped to clarify the elaborate signaling networks and the sophisticated crosstalk occurring among the different hormone signaling pathways. In this review, we summarize the roles of the major plant hormones in regulating abiotic and biotic stress responses with special focus on the significance of crosstalk between different hormones in generating a sophisticated and efficient stress response. We divided the discussion into the roles of ABA, salicylic acid, jasmonates and ethylene separately at the start of the review. Subsequently, we have discussed the crosstalk among them, followed by crosstalk with growth promoting hormones (gibberellins, auxins and cytokinins). These have been illustrated with examples drawn from selected abiotic and biotic stress responses. The discussion on seed dormancy and germination serves to illustrate the fine balance that can be enforced by the two key hormones ABA and GA in regulating plant responses to environmental signals. Conclusions The intricate web of crosstalk among the often redundant multitudes of signaling intermediates is just beginning to be understood. Future research employing genome-scale systems biology approaches to solve problems of such magnitude will undoubtedly lead to a better understanding of plant development. Therefore, discovering additional crosstalk mechanisms among various hormones in coordinating growth under stress will be an important theme in the field of abiotic stress research. Such efforts will help to reveal important points of genetic control that can be useful to engineer stress tolerant crops.
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                Author and article information

                Contributors
                Journal
                Front Plant Sci
                Front Plant Sci
                Front. Plant Sci.
                Frontiers in Plant Science
                Frontiers Media S.A.
                1664-462X
                12 April 2021
                2021
                : 12
                : 598173
                Affiliations
                [1] 1Biotechnology Research Institute, Heilongjiang Academy of Agricultural Sciences , Harbin, China
                [2] 2Biotechnology Research Institute, Chinese Academy of Agricultural Sciences , Beijing, China
                [3] 3National Key Facility of Crop Gene Resources and Genetic Improvement , Beijing, China
                Author notes

                Edited by: Xiaojuan Li, Beijing Forestry University, China

                Reviewed by: William Laing, The New Zealand Institute for Plant and Food Research Ltd., New Zealand; Tse-Min Lee, National Sun Yat-sen University, Taiwan

                *Correspondence: Zhijin Zhang, zhangzhijin@ 123456caas.cn

                These authors have contributed equally to this work

                This article was submitted to Plant Abiotic Stress, a section of the journal Frontiers in Plant Science

                Article
                10.3389/fpls.2021.598173
                8072462
                33912200
                1cf3a4ea-0ead-4fac-b3ad-f5e04c24d87f
                Copyright © 2021 Xiao, Li, Zhu, Wang, Zhang, Zheng, Zhao, Zhang, Wang and Zhang.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 24 August 2020
                : 17 March 2021
                Page count
                Figures: 3, Tables: 0, Equations: 0, References: 190, Pages: 13, Words: 12429
                Funding
                Funded by: National Basic Research Program of China 10.13039/501100012166
                Award ID: 31971838
                Award ID: 31670304
                Funded by: National Transgenic Major Program
                Award ID: 2018ZX08001-002
                Funded by: Central Non-profit Scientific Institution
                Award ID: Y2017PT25
                Funded by: National Key Research and Development Program of China 10.13039/501100012166
                Award ID: 2018YFD1000702
                Funded by: The Agricultural Science and Technology Innovation Program (ASTIP)
                Categories
                Plant Science
                Review

                Plant science & Botany
                ascorbate,abiotic stress,antioxidant,redox signal,phytohormone
                Plant science & Botany
                ascorbate, abiotic stress, antioxidant, redox signal, phytohormone

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