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.

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Metal accumulation in seeds is a prerequisite for germination and establishment of plants but also for micronutrient delivery to humans. To investigate metal transport processes and their interactions in seeds, we focused on METAL TOLERANCE PROTEIN8 (MTP8), a tonoplast transporter of the manganese (Mn) subclade of cation diffusion facilitators, which in Arabidopsis () is expressed in embryos of seeds. The x-ray fluorescence imaging showed that expression of MTP8 was responsible for Mn localization in subepidermal cells on the abaxial side of the cotyledons and in cortical cells of the hypocotyl. Accordingly, under low Mn availability, MTP8 increased seed stores of Mn, required for efficient seed germination. In mutant embryos lacking expression of (), MTP8 built up iron (Fe) hotspots in -expressing cells types, suggesting that MTP8 transports Fe in addition to Mn. In double mutant seeds, Mn and Fe were distributed in all cell types of the embryo. An Fe transport function of MTP8 was confirmed by its ability to complement Fe hypersensitivity of a yeast mutant defective in vacuolar Fe transport. Imbibing mutant seeds in the presence of Mn or subjecting seeds to wet-dry cycles showed that MTP8 conferred Mn tolerance. During germination, MTP8 promoted reallocation of Fe from the vasculature. These results indicate that cell type-specific accumulation of Mn and Fe in seeds depends on MTP8 and that this transporter plays an important role in the generation of seed metal stores as well as for metal homeostasis and germination efficiency under challenging environmental conditions.

The photosynthetic machinery of plants must be regulated to maximize the efficiency of light reactions and CO fixation. Changes in free Ca in the stroma of chloroplasts have been observed at the transition between light and darkness, and also in response to stress stimuli. Such Ca dynamics have been proposed to regulate photosynthetic capacity. However, the molecular mechanisms of Ca fluxes in the chloroplasts have been unknown. By employing a Ca reporter-based approach, we identified two chloroplast-localized Ca transporters in Arabidopsis thaliana, BICAT1 and BICAT2, that determine the amplitude of the darkness-induced Ca signal in the chloroplast stroma. BICAT2 mediated Ca uptake across the chloroplast envelope, and its knockout mutation strongly dampened the dark-induced [Ca ] signal. Conversely, this Ca transient was increased in knockout mutants of BICAT1, which transports Ca into the thylakoid lumen. Knockout mutation of BICAT2 caused severe defects in chloroplast morphology, pigmentation and photosynthetic light reactions, rendering bicat2 mutants barely viable under autotrophic growth conditions, while bicat1 mutants were less affected. These results show that BICAT transporters play a role in chloroplast Ca homeostasis. They are also involved in the regulation of photosynthesis and plant productivity. Further work will be required to reveal whether the effect on photosynthesis is a direct result of their role as Ca transporters.

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Background Plants are continuously exposed to changing environmental conditions and biotic attacks that affect plant growth. In crops, the inability to respond appropriately to stress has strong detrimental effects on agricultural production and yield. Ca 2+ signalling plays a fundamental role in the response of plants to most abiotic and biotic stresses. However, research on stimulus-specific Ca 2+ signals has mostly been pursued in Arabidopsis thaliana , while in other species these events are little investigated . Results In this study, we introduced the Ca 2+ reporter-encoding gene APOAEQUORIN into the crop species barley ( Hordeum vulgare ). Measurements of the dynamic changes in [Ca 2+ ] cyt in response to various stimuli such as NaCl, mannitol, H 2 O 2 , and flagellin 22 (flg22) revealed the occurrence of dose- as well as tissue-dependent [Ca 2+ ] cyt transients. Moreover, the Ca 2+ signatures were unique for each stimulus, suggesting the involvement of different Ca 2+ signalling components in the corresponding stress response. Alongside, the barley Ca 2+ signatures were compared to those produced by the phylogenetically distant model plant Arabidopsis. Notable differences in temporal kinetics and dose responses were observed, implying species-specific differences in stress response mechanisms. The plasma membrane Ca 2+ channel blocker La 3+ strongly inhibited the [Ca 2+ ] cyt response to all tested stimuli, indicating a critical role of extracellular Ca 2+ in the induction of stress-associated Ca 2+ signatures in barley. Moreover, by analysing spatio-temporal dynamics of the [Ca 2+ ] cyt transients along the developmental gradient of the barley leaf blade we demonstrate that different parts of the barley leaf show quantitative differences in [Ca 2+ ] cyt transients in response to NaCl and H 2 O 2 . There were only marginal differences in the response to flg22, indicative of developmental stage-dependent Ca 2+ responses specifically to NaCl and H 2 O 2 . Conclusion This study reveals tissue-specific Ca 2+ signals with stimulus-specific kinetics in the crop species barley, as well as quantitative differences al...

Manganese (Mn), iron (Fe), and zinc (Zn) are essential for diverse processes in plants, but their availability is often limiting or excessive. Cation diffusion facilitator (CDF) proteins have been implicated in the allocation of those metals in plants, whereby most of our mechanistic understanding has been obtained in Arabidopsis. It is unclear to what extent this can be generalized to other dicots. We characterized all CDFs/metal tolerance proteins of sugar beet (Beta vulgaris spp. vulgaris), which is phylogenetically distant from Arabidopsis. Analysis of subcellular localization, substrate selectivities, and transcriptional regulation upon exposure to metal deficiencies and toxicities revealed unexpected deviations from their Arabidopsis counterparts. Localization and selectivity of some members were modulated by alternative splicing. Notably, unlike in Arabidopsis, Mn-and Zn-sequestrating members were not induced in Fe-deficient roots, pointing to differences in the Fe acquisition machinery. This was supported by low Zn and Mn accumulation under Fe deficiency and a strikingly increased Fe accumulation under Mn and Zn excess, coinciding with an induction of BvIRT1. High Zn load caused a massive upregulation of Zn-BvMTPs. The results suggest that the employment of the CDF toolbox is highly diverse amongst dicots, which questions the general applicability of metal homeostasis models derived from Arabidopsis.

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