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      Reducing Cadmium Accumulation in Plants: Structure–Function Relations and Tissue-Specific Operation of Transporters in the Spotlight

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          Abstract

          Cadmium (Cd) is present in many soils and, when entering the food chain, represents a major health threat to humans. Reducing Cd accumulation in plants is complicated by the fact that most known Cd transporters also operate in the transport of essential nutrients such as Zn, Fe, Mn, or Cu. This work summarizes the current knowledge of mechanisms mediating Cd uptake, radial transport, and translocation within the plant. It is concluded that real progress in the field may be only achieved if the transport of Cd and the above beneficial micronutrients is uncoupled, and we discuss the possible ways of achieving this goal. Accordingly, we suggest that the major focus of research in the field should be on the structure–function relations of various transporter isoforms and the functional assessment of their tissue-specific operation. Of specific importance are two tissues. The first one is a xylem parenchyma in plant roots; a major “controller” of Cd loading into the xylem and its transport to the shoot. The second one is a phloem tissue that operates in the last step of a metal transport. Another promising and currently underexplored avenue is to understand the role of non-selective cation channels in Cd uptake and reveal mechanisms of their regulation.

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          Most cited references135

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          Toxic metal accumulation, responses to exposure and mechanisms of tolerance in plants.

          S Clemens (2006)
          Over the past 200 years emissions of toxic heavy metals have risen tremendously and significantly exceed those from natural sources for practically all metals. Uptake and accumulation by crop plants represents the main entry pathway for potentially health-threatening toxic metals into human and animal food. Of major concern are the metalloids arsenic (As) and selenium (Se), and the metals cadmium (Cd), mercury (Hg), and lead (Pb). This review discusses the molecular mechanisms of toxic metal accumulation in plants and algae, the responses to metal exposure, as well as our understanding of metal tolerance and its evolution. The main emphasis will be on cadmium, which is by far the most widely studied of the non-essential toxic metals/metalloids. Entry via Zn2+, Fe2+, and Ca2+ transporters is the molecular basis of Cd2+ uptake into plant cells. Much less is known about the partitioning of non-essential metals and about the genes underlying the enormous diversity among plants with respect to Cd accumulation in different tissues. Numerous studies have described symptoms and responses of plants upon toxic metal exposure. Mysterious are primary targets of toxicity, the degree of specificity of responses, the sensing and the signaling events that lead to transcriptional activation. All plants apparently possess a basal tolerance of toxic non-essential metals. For Cd and As, this is largely dependent on the phytochelatin pathway. Not understood is the molecular biology of Cd hypertolerance in certain plant species such as the metallophytes Arabidopsis halleri or Thlaspi caerulescens.
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            Nramp5 is a major transporter responsible for manganese and cadmium uptake in rice.

            Paddy rice (Oryza sativa) is able to accumulate high concentrations of Mn without showing toxicity; however, the molecular mechanisms underlying Mn uptake are unknown. Here, we report that a member of the Nramp (for the Natural Resistance-Associated Macrophage Protein) family, Nramp5, is involved in Mn uptake and subsequently the accumulation of high concentrations of Mn in rice. Nramp5 was constitutively expressed in the roots and encodes a plasma membrane-localized protein. Nramp5 was polarly localized at the distal side of both exodermis and endodermis cells. Knockout of Nramp5 resulted in a significant reduction in growth and grain yield, especially when grown at low Mn concentrations. This growth reduction could be partially rescued by supplying high concentrations of Mn but not by the addition of Fe. Mineral analysis showed that the concentration of Mn and Cd in both the roots and shoots was lower in the knockout line than in wild-type rice. A short-term uptake experiment revealed that the knockout line lost the ability to take up Mn and Cd. Taken together, Nramp5 is a major transporter of Mn and Cd and is responsible for the transport of Mn and Cd from the external solution to root cells.
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              Toxic Heavy Metal and Metalloid Accumulation in Crop Plants and Foods.

              Arsenic, cadmium, lead, and mercury are toxic elements that are almost ubiquitously present at low levels in the environment because of anthropogenic influences. Dietary intake of plant-derived food represents a major fraction of potentially health-threatening human exposure, especially to arsenic and cadmium. In the interest of better food safety, it is important to reduce toxic element accumulation in crops. A molecular understanding of the pathways responsible for this accumulation can enable the development of crop varieties with strongly reduced concentrations of toxic elements in their edible parts. Such understanding is rapidly progressing for arsenic and cadmium but is in its infancy for lead and mercury. Basic discoveries have been made in Arabidopsis, rice, and other models, and most advances in crops have been made in rice. Proteins mediating the uptake of arsenic and cadmium have been identified, and the speciation and biotransformations of arsenic are now understood. Factors controlling the efficiency of root-to-shoot translocation and the partitioning of toxic elements through the rice node have also been identified.
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                Author and article information

                Journal
                Plants (Basel)
                Plants (Basel)
                plants
                Plants
                MDPI
                2223-7747
                09 February 2020
                February 2020
                : 9
                : 2
                : 223
                Affiliations
                [1 ]International Research Center for Environmental Membrane Biology, Foshan University, Foshan 528000, China; Xin.Huang@ 123456utas.edu.au (X.H.); songpo.duan@ 123456hotmail.com (S.D.); qi.wu@ 123456fosu.edu.cn (Q.W.); yumin@ 123456fosu.edu.cn (M.Y.)
                [2 ]Tasmanian Institute of Agriculture, University of Tasmania, Hobart TAS 7001, Australia
                Author notes
                Article
                plants-09-00223
                10.3390/plants9020223
                7076666
                32050442
                1c511d3a-e1aa-4c60-a64b-e65d4fbf5fd7
                © 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
                : 31 January 2020
                : 06 February 2020
                Categories
                Review

                cadmium toxicity,membrane transport,tissue tolerance,nramp,irt,mtp,non-selective cation channel

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