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      Copper Trafficking in Plants and Its Implication on Cell Wall Dynamics

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          Abstract

          In plants, copper (Cu) acts as essential cofactor of numerous proteins. While the definitive number of these so-called cuproproteins is unknown, they perform central functions in plant cells. As micronutrient, a minimal amount of Cu is needed to ensure cellular functions. However, Cu excess may exert in contrast detrimental effects on plant primary production and even survival. Therefore it is essential for a plant to have a strictly controlled Cu homeostasis, an equilibrium that is both tissue and developmentally influenced. In the current review an overview is presented on the different stages of Cu transport from the soil into the plant and throughout the different plant tissues. Special emphasis is on the Cu-dependent responses mediated by the SPL7 transcription factor, and the crosstalk between this transcriptional regulation and microRNA-mediated suppression of translation of seemingly non-essential cuproproteins. Since Cu is an essential player in electron transport, we also review the recent insights into the molecular mechanisms controlling chloroplastic and mitochondrial Cu transport and homeostasis. We finally highlight the involvement of numerous Cu-proteins and Cu-dependent activities in the properties of one of the major Cu-accumulation sites in plants: the cell wall.

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          Physiological functions of mineral micronutrients (Cu, Zn, Mn, Fe, Ni, Mo, B, Cl).

          Micronutrients are involved in all metabolic and cellular functions. Plants differ in their need for micronutrients, and we will focus here only on those elements that are generally accepted as essential for all higher plants: boron (B), chloride (Cl), copper (Cu), iron (Fe), manganese (Mn), molybdenum (Mo), nickel (Ni), and zinc (Zn). Several of these elements are redox-active that makes them essential as catalytically active cofactors in enzymes, others have enzyme-activating functions, and yet others fulfill a structural role in stabilizing proteins. In this review, we focus on the major functions of mineral micronutrients, mostly in cases where they were shown as constituents of proteins, making a selection and highlighting some functions in more detail.
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            Ethylene: a gaseous signal molecule in plants.

            Ethylene regulates a multitude of plant processes, ranging from seed germination to organ senescence. Of particular economic importance is the role of ethylene as an inducer of fruit ripening. Ethylene is synthesized from S-adenosyl-L-methionine via 1-aminocyclopropane-1-carboxylic acid (ACC). The enzymes catalyzing the two reactions in this pathway are ACC synthase and ACC oxidase. Environmental and endogenous signals regulate ethylene biosynthesis primarily through differential expression of ACC synthase genes. Components of the ethylene signal transduction pathway have been identified by characterization of ethylene-response mutants in Arabidopsis thaliana. One class of mutations, exemplified by etr1, led to the identification of the ethylene receptors, which turned out to be related to bacterial two-component signaling systems. Mutations that eliminate ethylene binding to the receptor yield a dominant, ethylene-insensitive phenotype. CTR1 encodes a Raf-like Ser/Thr protein kinase that acts downstream from the ethylene receptor and may be part of a MAP kinase cascade. Mutants in CTR1 exhibit a constitutive ethylene-response phenotype. Both the ethylene receptors and CTR1 are negative regulators of ethylene responses. EIN2 and EIN3 are epistatic to CTR1, and mutations in either gene lead to ethylene insensitivity. Whereas the function of EIN2 in ethylene transduction is not known, EIN3 is a putative transcription factor involved in regulating expression of ethylene-responsive genes. Biotechnological modifications of ethylene synthesis and of sensitivity to ethylene are promising methods to prevent spoilage of agricultural products such as fruits, whose ripening is induced by ethylene.
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              The essential basic helix-loop-helix protein FIT1 is required for the iron deficiency response.

              Regulation of iron uptake is critical for plant survival. Although the activities responsible for reduction and transport of iron at the plant root surface have been described, the genes controlling these activities are largely unknown. We report the identification of the essential gene Fe-deficiency Induced Transcription Factor 1 (FIT1), which encodes a putative transcription factor that regulates iron uptake responses in Arabidopsis thaliana. Like the Fe(III) chelate reductase FRO2 and high affinity Fe(II) transporter IRT1, FIT1 mRNA is detected in the outer cell layers of the root and accumulates in response to iron deficiency. fit1 mutant plants are chlorotic and die as seedlings but can be rescued by the addition of supplemental iron, pointing to a defect in iron uptake. fit1 mutant plants accumulate less iron than wild-type plants in root and shoot tissues. Microarray analysis shows that expression of many (72 of 179) iron-regulated genes is dependent on FIT1. We demonstrate that FIT1 regulates FRO2 at the level of mRNA accumulation and IRT1 at the level of protein accumulation. We propose a new model for iron uptake in Arabidopsis where FRO2 and IRT1 are differentially regulated by FIT1.
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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
                06 May 2016
                2016
                : 7
                : 601
                Affiliations
                [1] 1Environmental Research and Innovation Department, Luxembourg Institute of Science and Technology Esch-sur-Alzette, Luxembourg
                [2] 2Groupe de Recherche en Physiologie Végétale, Earth and Life Institute Agronomy, Université catholique de Louvain Louvain-la-Neuve, Belgium
                Author notes

                Edited by: Jon Pittman, University of Manchester, UK

                Reviewed by: Olena Vatamaniuk, Cornell University, USA; Sergi Puig, Instituto de Agroquimica y Tecnologia de Alimentos-Consejo Superior de Investigaciones Científicas, Spain

                *Correspondence: Kjell Sergeant, kjell.sergeant@ 123456list.lu

                This article was submitted to Plant Traffic and Transport, a section of the journal Frontiers in Plant Science

                Article
                10.3389/fpls.2016.00601
                4859090
                27200069
                8150efaf-bbdf-4e21-af75-9d11a440883d
                Copyright © 2016 Printz, Lutts, Hausman and Sergeant.

                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
                : 17 November 2015
                : 18 April 2016
                Page count
                Figures: 4, Tables: 0, Equations: 0, References: 129, Pages: 16, Words: 0
                Funding
                Funded by: Fonds National de la Recherche Luxembourg 10.13039/501100001866
                Award ID: C13/SR/5774202
                Categories
                Plant Science
                Review

                Plant science & Botany
                copper,transport,plant,cell wall
                Plant science & Botany
                copper, transport, plant, cell wall

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