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      Flavonoids: a metabolic network mediating plants adaptation to their real estate

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          Abstract

          From an evolutionary perspective, the emergence of the sophisticated chemical scaffolds of flavonoid molecules represents a key step in the colonization of Earth’s terrestrial environment by vascular plants nearly 500 million years ago. The subsequent evolution of flavonoids through recruitment and modification of ancestors involved in primary metabolism has allowed vascular plants to cope with pathogen invasion and damaging UV light. The functional properties of flavonoids as a unique combination of different classes of compounds vary significantly depending on the demands of their local real estate. Apart from geographical location, the composition of flavonoids is largely dependent on the plant species, their developmental stage, tissue type, subcellular localization, and key ecological influences of both biotic and abiotic origin. Molecular and metabolic cross-talk between flavonoid and other pathways as a result of the re-direction of intermediate molecules have been well investigated. This metabolic plasticity is a key factor in plant adaptive strength and is of paramount importance for early land plants adaptation to their local ecosystems. In human and animal health the biological and pharmacological activities of flavonoids have been investigated in great depth and have shown a wide range of anti-inflammatory, anti-oxidant, anti-microbial, and anti-cancer properties. In this paper we review the application of advanced gene technologies for targeted reprogramming of the flavonoid pathway in plants to understand its molecular functions and explore opportunities for major improvements in forage plants enhancing animal health and production.

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          Flavonoids: biosynthesis, biological functions, and biotechnological applications

          Flavonoids are widely distributed secondary metabolites with different metabolic functions in plants. The elucidation of the biosynthetic pathways, as well as their regulation by MYB, basic helix-loop-helix (bHLH), and WD40-type transcription factors, has allowed metabolic engineering of plants through the manipulation of the different final products with valuable applications. The present review describes the regulation of flavonoid biosynthesis, as well as the biological functions of flavonoids in plants, such as in defense against UV-B radiation and pathogen infection, nodulation, and pollen fertility. In addition, we discuss different strategies and achievements through the genetic engineering of flavonoid biosynthesis with implication in the industry and the combinatorial biosynthesis in microorganisms by the reconstruction of the pathway to obtain high amounts of specific compounds.
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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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              Recruitment of entomopathogenic nematodes by insect-damaged maize roots.

              Plants under attack by arthropod herbivores often emit volatile compounds from their leaves that attract natural enemies of the herbivores. Here we report the first identification of an insect-induced belowground plant signal, (E)-beta-caryophyllene, which strongly attracts an entomopathogenic nematode. Maize roots release this sesquiterpene in response to feeding by larvae of the beetle Diabrotica virgifera virgifera, a maize pest that is currently invading Europe. Most North American maize lines do not release (E)-beta-caryophyllene, whereas European lines and the wild maize ancestor, teosinte, readily do so in response to D. v. virgifera attack. This difference was consistent with striking differences in the attractiveness of representative lines in the laboratory. Field experiments showed a fivefold higher nematode infection rate of D. v. virgifera larvae on a maize variety that produces the signal than on a variety that does not, whereas spiking the soil near the latter variety with authentic (E)-beta-caryophyllene decreased the emergence of adult D. v. virgifera to less than half. North American maize lines must have lost the signal during the breeding process. Development of new varieties that release the attractant in adequate amounts should help enhance the efficacy of nematodes as biological control agents against root pests like D. v. virgifera.
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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
                28 July 2014
                10 November 2014
                2014
                : 5
                : 620
                Affiliations
                [1] 1Royal Melbourne Institute of Technology University Bundoora, VIC, Australia
                [2] 2Department of Environment and Primary Industries, Biosciences Research Division, AgriBio, Centre for AgriBioscience Bundoora, VIC, Australia
                [3] 3School of Applied Systems Biology, La Trobe University – AgriBio, Centre for AgriBioscience Bundoora, VIC, Australia
                Author notes

                Edited by: Francesco Paolocci, Italian National Research Council – Institute of Plant Genetics Division of Perugia, Italy

                Reviewed by: Akifumi Sugiyama, Kyoto University, Japan; Marcello Iriti, Milan State University, Italy

                *Correspondence: German Spangenberg, Department of Environment and Primary Industries, Biosciences Research Division, AgriBio, Centre for AgriBioscience – School of Applied Systems Biology, La Trobe University, 5 Ring Road, Bundoora, VIC 3083, Australia e-mail: german.spangenberg@ 123456depi.vic.gov.au

                This article was submitted to Plant Metabolism and Chemodiversity, a section of the journal Frontiers in Plant Science.

                Article
                10.3389/fpls.2014.00620
                4226159
                25426130
                acbb5edd-b20a-44d6-a68f-89821445f8da
                Copyright © 2014 Mouradov and Spangenberg.

                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) or licensor 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
                : 02 July 2014
                : 21 October 2014
                Page count
                Figures: 6, Tables: 0, Equations: 0, References: 169, Pages: 16, Words: 0
                Categories
                Plant Science
                Review Article

                Plant science & Botany
                flavonoids,anthocyanins,proanthocyanidins,evolution,transgenic,metabolism
                Plant science & Botany
                flavonoids, anthocyanins, proanthocyanidins, evolution, transgenic, metabolism

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