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      Root exudate metabolomes change under drought and show limited capacity for recovery

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          Abstract

          Root exudates comprise a large variety of compounds released by plants into the rhizosphere, including low-molecular-weight primary metabolites (particularly saccharides, amino acids and organic acids) and secondary metabolites (phenolics, flavonoids and terpenoids). Changes in exudate composition could have impacts on the plant itself, on other plants, on soil properties (e.g. amount of soil organic matter), and on soil organisms. The effects of drought on the composition of root exudates, however, have been rarely studied. We used an ecometabolomics approach to identify the compounds in the exudates of Quercus ilex (holm oak) under an experimental drought gradient and subsequent recovery. Increasing drought stress strongly affected the composition of the exudate metabolome. Plant exudates under drought consisted mainly of secondary metabolites (71% of total metabolites) associated with plant responses to drought stress, whereas the metabolite composition under recovery shifted towards a dominance of primary metabolites (81% of total metabolites). These results strongly suggested that roots exude the most abundant root metabolites. The exudates were changed irreversibly by the lack of water under extreme drought conditions, and the plants could not recover.

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          Metabolomics--the link between genotypes and phenotypes.

          Metabolites are the end products of cellular regulatory processes, and their levels can be regarded as the ultimate response of biological systems to genetic or environmental changes. In parallel to the terms 'transcriptome' and proteome', the set of metabolites synthesized by a biological system constitute its 'metabolome'. Yet, unlike other functional genomics approaches, the unbiased simultaneous identification and quantification of plant metabolomes has been largely neglected. Until recently, most analyses were restricted to profiling selected classes of compounds, or to fingerprinting metabolic changes without sufficient analytical resolution to determine metabolite levels and identities individually. As a prerequisite for metabolomic analysis, careful consideration of the methods employed for tissue extraction, sample preparation, data acquisition, and data mining must be taken. In this review, the differences among metabolite target analysis, metabolite profiling, and metabolic fingerprinting are clarified, and terms are defined. Current approaches are examined, and potential applications are summarized with a special emphasis on data mining and mathematical modelling of metabolism.
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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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              Regulation and function of root exudates

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                Author and article information

                Contributors
                albert.gargallo@gmail.com
                Journal
                Sci Rep
                Sci Rep
                Scientific Reports
                Nature Publishing Group UK (London )
                2045-2322
                23 August 2018
                23 August 2018
                2018
                : 8
                : 12696
                Affiliations
                [1 ]ISNI 0000 0001 2183 4846, GRID grid.4711.3, CSIC, Global Ecology Unit CREAF- CSIC-UAB, ; Bellaterra, 08193 Catalonia Spain
                [2 ]ISNI 0000 0001 0722 403X, GRID grid.452388.0, CREAF, Cerdanyola del Vallès, ; 08193 Catalonia, Spain
                [3 ]ISNI 0000 0001 1015 3316, GRID grid.418095.1, Global Change Research Institute, The Czech Academy of Sciences, ; Belidla 986/4a, CZ-60300 Brno, Czech Republic
                Author information
                http://orcid.org/0000-0001-6584-3541
                http://orcid.org/0000-0002-7215-0150
                Article
                30150
                10.1038/s41598-018-30150-0
                6107494
                30140025
                5dd3d5c5-679c-4933-9138-0608dc3f4c31
                © The Author(s) 2018

                Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

                History
                : 6 March 2018
                : 10 July 2018
                Funding
                Funded by: FundRef https://doi.org/10.13039/501100000781, EC | European Research Council (ERC);
                Award ID: ERC-2013-SyG-610028
                Award ID: ERC-2013-SyG-610028
                Award ID: ERC-2013-SyG-610028
                Award ID: ERC-2013-SyG-610028
                Award Recipient :
                Funded by: FundRef https://doi.org/10.13039/501100001624, European Council of International Schools (ECIS);
                Award ID: ERC-2013-SyG-610028
                Award Recipient :
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