Manganese (Mn) is an important micronutrient for plant growth and development and sustains metabolic roles within different plant cell compartments. The metal is an essential cofactor for the oxygen-evolving complex (OEC) of the photosynthetic machinery, catalyzing the water-splitting reaction in photosystem II (PSII). Despite the importance of Mn for photosynthesis and other processes, the physiological relevance of Mn uptake and compartmentation in plants has been underrated. The subcellular Mn homeostasis to maintain compartmented Mn-dependent metabolic processes like glycosylation, ROS scavenging, and photosynthesis is mediated by a multitude of transport proteins from diverse gene families. However, Mn homeostasis may be disturbed under suboptimal or excessive Mn availability. Mn deficiency is a serious, widespread plant nutritional disorder in dry, well-aerated and calcareous soils, as well as in soils containing high amounts of organic matter, where bio-availability of Mn can decrease far below the level that is required for normal plant growth. By contrast, Mn toxicity occurs on poorly drained and acidic soils in which high amounts of Mn are rendered available. Consequently, plants have evolved mechanisms to tightly regulate Mn uptake, trafficking, and storage. This review provides a comprehensive overview, with a focus on recent advances, on the multiple functions of transporters involved in Mn homeostasis, as well as their regulatory mechanisms in the plant's response to different conditions of Mn availability.

/pdf/manganese-in-plants-from-acquisition-to-subcellular-1vn9c3muvz.pdf

Manganese in Plants: From Acquisition to Subcellular Allocation.

Metal homeostasis is critical for plant growth, development and adaptation to environmental stresses and largely governed by a variety of metal transporters. The plant ZIP (Zn-regulated transporter, Iron-regulated transporter-like Protein) family proteins belong to the integral membrane transporters responsible for uptake and allocation of essential and non-essential metals. However, whether the ZIP family members mediate metal efflux and its regulatory mechanism remains unknown. In this report, we provided evidence that OsZIP1 is a metal-detoxified transporter through preventing excess Zn, Cu and Cd accumulation in rice. OsZIP1 is abundantly expressed in roots throughout the life span and sufficiently induced by excess Zn, Cu and Cd but not by Mn and Fe at transcriptional and translational levels. Expression of OsZIP-GFP fusion in rice protoplasts and tobacco leaves shows that OsZIP1 resides in the endoplasmic reticulum (ER) and plasma membrane (PM). The yeast (Saccharomyces cerevisiae) complementation test shows that expression of OsZIP1 reduced Zn accumulation. Transgenic rice overexpressing OsZIP1 grew better under excess metal stress but accumulated less of the metals in plants. In contrast, both oszip1 mutant and RNA interference (RNAi) lines accumulated more metal in roots and contributed to metal sensitive phenotypes. These results suggest OsZIP1 is able to function as a metal exporter in rice when Zn, Cu and Cd are excess in environment. We further identified the DNA methylation of histone H3K9me2 of OsZIP1 and found that OsZIP1 locus, whose transcribed regions imbed a 242 bp sequence, is demethylated, suggesting that epigenetic modification is likely associated with OsZIP1 function under Cd stress. OsZIP1 is a transporter that is required for detoxification of excess Zn, Cu and Cd in rice.

/pdf/oszip1-functions-as-a-metal-efflux-transporter-limiting-5bqjynoz8l.pdf

OsZIP1 functions as a metal efflux transporter limiting excess zinc, copper and cadmium accumulation in rice

Crop diseases threaten food security and sustainable agriculture. Consumption of crops containing non-essential toxic metals leads to health risks for humans. Therefore, breeding disease-resistance and toxic metal-safe crops are a double gain strategy contributing to food security. Here, we show that rice heavy metal transporter OsNRAMP1 plays important roles in plant immunity by modulating metal ions and reactive oxygen species (ROS) homeostasis. OsNRAMP1 expression was induced after pathogenic bacteria and fungi infection. The osnramp1 mutants had increased content of H2 O2 and activity of superoxide dismutase, but decreased activity of catalase, exhibiting enhanced broad-spectrum resistance against bacterial and fungal pathogens. RNA-seq analysis identified a number of differentially expressed genes that were involved in metal ions and ROS homeostasis. Altered expression of metal ion-dependent ROS-scavenging enzymes genes and lower accumulation of cations such as Mn coordinately triggered compromised metal ion-dependent enzyme-catalyzing activity and modulated ROS homeostasis, which together contributed to disease resistance in osnramp1 mutants. Furthermore, the osnramp1 mutants contained lower levels of toxic heavy metals Cd and Pb and micronutrients Ni and Mn in leaves and grains. Taken together, a proof-of-concept was achieved that broad-spectrum disease resistance and toxic heavy metal-safe rice was engineered by removal of OsNRAMP1 gene. This article is protected by copyright. All rights reserved.

The rice heavy metal transporter OsNRAMP1 regulates disease resistance by modulating ROS homeostasis.

\n Calcium (Ca2+) and manganese (Mn2+) are essential elements for plants and have similar ionic radii and binding coordination. They are assigned specific functions within organelles, but share many transport mechanisms to cross organellar membranes. Despite their points of interaction, those elements are usually investigated and reviewed separately. This review takes them out of this isolation. It highlights our current mechanistic understanding and points to open questions of their functions, their transport, and their interplay in the endoplasmic reticulum (ER), vesicular compartments [Golgi apparatus, trans-Golgi Network (TGN), prevacuolar compartment (PVC)], vacuoles, chloroplasts, mitochondria, and peroxisomes. Complex processes demanding these cations, such as Mn2+-dependent glycosylation or systemic Ca2+ signaling, are covered in some detail if they have not been reviewed recently or if recent findings add to current models. The function of Ca2+ as signaling agent released from organelles into the cytosol and within the organelles themselves is a recurrent theme of this review, again keeping the interference by Mn2+ in mind. The involvement of organellar channels [e.g., Glutamate-Receptor-Likes (GLRs), Cyclic-Nucleotide-Gated Channels (CNGCs), Mitochondrial Conductivity Units (MCUs), Two-Pore Channel1 (TPC1)], transporters [e.g., Natural-Resistance-Associated Macrophage Proteins (NRAMPs), Calcium Exchangers (CAXs), Metal-Tolerance Proteins (MTPs), Bivalent-Cation Transporters (BICATs)] and pumps [Autoinhibited Ca2+-ATPases (ACAs), ER Ca2+-ATPases (ECAs)] in the import and export of organellar Ca2+ and Mn2+ is scrutinized, whereby current controversial issues are pointed out. Mechanisms in animals and yeast are taken into account where they may provide a blueprint for processes in plants, in particular with respect to tunable molecular mechanisms of Ca2+-versus-Mn2+ selectivity.

/pdf/transport-functions-and-interaction-of-calcium-and-manganese-3ld0895e79.pdf

Transport, functions, and interaction of calcium and manganese in plant organellar compartments

Toxicity of copper hydroxide nanoparticles, bulk copper hydroxide, and ionic copper to alfalfa plants: A spectroscopic and gene expression study.

OsMTP11, a trans-Golgi network localized transporter, is involved in manganese tolerance in rice.

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OsMTP11, a trans-Golgi network localized transporter, is involved in manganese tolerance in rice.