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Characterization of Arabidopsis COPT family members. (a) Subcellular localization in Arabidopsis protoplasts. COPT-GFP protein localization is indicated in green. (b) Transcript regulation in response to copper deficiency. (c) Expression pattern in flowers. (d) Expression pattern in roots and shoots. (e) Parameters affected in copt mutant lines upon copper scarcity. COPT3 and COPT4 have not been included due to the small amount of data currently available.

Characterization of Arabidopsis COPT family members. (a) Subcellular localization in Arabidopsis protoplasts. COPT-GFP protein localization is indicated in green. (b) Transcript regulation in response to copper deficiency. (c) Expression pattern in flowers. (d) Expression pattern in roots and shoots. (e) Parameters affected in copt mutant lines upon copper scarcity. COPT3 and COPT4 have not been included due to the small amount of data currently available.

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Copper (Cu) is an essential micronutrient for all eukaryotes because it participates as a redox active cofactor in multiple biological processes, including mitochondrial respiration, photosynthesis, oxidative stress protection, and iron (Fe) transport. In eukaryotic cells, Cu transport toward the cytoplasm is mediated by the conserved CTR/COPT fami...

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... Many transporters that mediate HM uptake in the cell have quite limited specificity, which is probably compensated for by the tight regulation of transporters at the transcriptional level [41]. The copper is likely reduced by Cu 2+ -reducing FRO from Cu 2+ to Cu + outside of the cell and then transported through the plasmalemma by copper transporters (COPTs), which are highly specific to monovalent Cu + ions, after which Cu+ is specifically delivered to intracellular targets by Cu chaperones [46], leaving free Cu + cytosolic concentrations at very low (likely attomolar) levels [47]. Copper uptake is likely regulated more tightly than that of other essential transition metals, such as Zn, Mn, or Fe, due to the greater ability of Cu to bind to intracellular sites and the deleterious effects of such binding on intracellular processes. ...
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... Micro-nutrients, such as divalent cations (Zn 2þ , Fe 2þ , and Cu 2þ ), share common transporters for plant acquisition and redistribution (Puig and Peñarrubia, 2009). However they have primary transporters, including the acquisition and export of Zn by the zinc-regulated, iron-regulated transporter-like proteins (ZIPs) (Ajeesh Krishna et al., 2020) and Heavy Metal ATPase2 (HMA2 and 4) transporters (Hanikenne et al., 2008;Qiao et al., 2018) respectively, Fe by Fe-regulated transporter 1 (IRT1) (Eide et al., 1996;Vert et al., 2002) and IRT1/FRDL transporters (Kobayashi et al., 2019), Mn by natural resistanceassociated macrophage proteins (NRAMP) and metal tolerance protein 9 (MTP9) transporters (Ehrnstorfer et al., 2017;Zhao Qiufang et al., 2019) Cu by Cu transporters (COPTs), NRAMP and HMA5 (Puig, 2014;Sanz et al., 2019;Kumar et al., 2021). ZIP also transports metal cations in addition to Zn 2þ such as manganese (Mn 2þ ), iron (Fe 2þ ), copper (Cu 2þ ), cobalt (Co 2þ ), nickel (Ni 2þ ), and nonnutrient metals like cadmium (Cd 2þ ) (Pedas and Husted, 2009). ...
Article
Plants require essential nutrients for maintaining normal physiological, biochemical, and molecular functions. Plasma membrane-localized nutrient transporters play key roles in acquiring and allocating plant nutrients. Extensive studies have been performed on the functional characterization of key plant nutrient transporters in the past decades. Crystal structures of a few plant nutrient transporters and bacterial or fungal homologs were solved, which helped to predict the key residues and transport mechanisms of plant nutrient transporters. Site-directed mutagenesis and yeast complementation studies have also identified functional residues. This review presents a comprehensive list of the functional residues of various macro-and micro-nutrient transporters involved in acquiring and redistributing nutrients in the plant. We have also analyzed the functionally important residues of various plant nutrient transporters with bioinformatics tools. We then draw insights on the possible application of CRISPR/Cas tools to edit key residues to improve nutrient transport and agronomical performance. Utilization of genome editing tools like CRISPR could help develop DNA-free GM crops for quicker approval for field cultivation, contributing to food security amidst global climate change.
... When available in growth media, Cu 2+ enters plant root cells through promiscuous divalent cation transporters (YSL, ZIP) [11,17,18]. However, when scarce, plants use a Cu + -specific mobilization system based on the acidification of the external medium by H + -ATPases, plasma membrane NADPH-dependent cupric reductases (FRO4 and FRO5) [13,19], and the participation of high-affinity Cu + transporters (COPTs) that incorporate Cu + inside cells from the apoplast [12,[20][21][22]. In line with this, there is a prevailing dogma of protein-protein interactions that mediate Cu + delivery by which COPT members facilitate Cu + entry across the membrane and modulate its transfer to specific metallochaperones for targeted distribution to different locations and organelles [23,24]. ...
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... COPT2, another Cu(I) transporter of the COPT/Ctr-like family is primarily localized in the epidermal cells of the root, root hairs but are absent in the meristematic zones and shows upregulation in response to Cu depletion in an SPL dependent way. It primarily mediates the secondary pathway of root Cu incorporation and is mainly present in the reproductive organs (Puig, 2014;Gayomba et al., 2013). ...
... In terrestrial environments, P compounds act as a source of heavy metals for plants either by direct metal adsorption, phosphate anion-induced metal adsorption and desorption or modification of the rhizosphere through acidification and mycorrhizal associations [82]. Copper is associated with its transporters from the surrounding medium to plant tissues, such as the COPT family of proteins, which also participate in the metabolism of phosphate [74,83,84]. Similar to P, N is also a macronutrient and is essential for plant growth [85,86]. ...
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Heavy metal pollution creates environmental health concerns. Among these, iron (Fe), copper (Cu) and manganese (Mn) are commonly found in aquatic environments due to the release of wastewaters. Phytoremediation in hydroponics uses macrophytes to treat contaminated environments, and this is influenced by environmental factors. However, the relationship between these factors and the removal of Fe, Cu and Mn by macrophytes is not known. Therefore, a meta-analysis serves to determine the correlations between environmental factors and the removal of these metals in real wastewater by macrophytes, as well as to identify the role of different aquatic forms of macrophytes in phytoremediation. Emergent macrophytes had higher concentrations of manganese in their tissues, and higher bioconcentrations factor of iron and manganese than floating plants. Regardless of the biotope, higher concentrations of Fe and Cu decreased the ability of plants to bioconcentrate them. The correlations among exposure time, pH, dissolved oxygen, nitrogen, phosphorus, photoperiod and metal phytoremediation by plants were also found. It can be concluded that the emergent macrophytes showed better performance in terms of the removal of Fe, Cu and Mn, and that the significant correlations between environmental factors and removal vary with the type of metal and the environmental factor analyzed.
... MDA levels, an indicator of lipid peroxidation due to action of ROS over cell membranes, were not affected by the respective alterations of H 2 O 2 levels, suggesting that Fe deprivation treatment was not severe enough to inflict significant cell damage. When Fe is scarce, there is an increased participation of other divalent cations, such as Cu, Zn, and Mn, as prosthetic metals in the active sites of ROS-scavenging enzymes (Waters and Armbrust 2013;Puig 2014). In line with this, the accumulation of Zn and Mn in the leaves of WT and hp1, and Cu in the roots of all genotypes in substitution of Fe in the composition of the antioxidant enzymes might have helped to contend ROS burst and consequent increased MDA levels. ...
Article
Iron (Fe) is a micronutrient for plant development, as constituent of several photosynthesis- and respiration-related proteins and enzymes. Consequently, Fe deficiency leads to chlorosis in leaves and plant growth impairment. It has become increasingly evident that light signals coordinate iron homeostasis in plants. To further address new insights into how light is a fundamental part of Fe deficiency responses, we employed Micro-Tom (wild type, WT) tomato (Solanum lycopersicum L.) plants and high-pigment 1 (hp1) and aurea (au) photomorphogenic mutants, which exhibit an excessive light response and low light perception, respectively. Plant growth, pigment contents, oxidative status, and nutrient profile were analyzed. The results revealed the influence of the different genotypes on Fe deficiency responses. WT and au exhibited plant growth reduction under Fe deficiency. WT, hp1 and au demonstrated that Fe availability and light perception play fundamental roles in chlorophyll and anthocyanin biosynthesis. Lipid peroxidation was not increased for any genotype under Fe deficiency, indicating that mutations in light perception and signaling differentially modulate H2O2 production and scavenging under this condition. Additionally, macronutrients and micronutrients were taken up and distributed differently among the different plant genotypes, tissues and Fe conditions analyzed. In general, the au plants accumulated lower amounts of nutrients (Ca, S, P, Mg, B and Zn) than the WT and hp1 genotypes regardless of the Fe concentrations. Our data clearly indicates that light perception and signaling influence Fe-dependent morphophysiological responses in plants, suggesting possibilities for biotechnological improvement of crops grown under Fe shortage.
... COPT proteins have been reported in transfer of Cu in many major parts of plants. Puig (2014) reviewed and stated that COPT1 plays role in absorption of Cu in plant roots, COPT6 plays role in the distribution of Cu in plant shoots, and COPT5 activates and organizes Cu from organelles meant for storage. Therefore, in the light of aforementioned functions of COPT proteins, it can be stated that the COPT proteins play an indispensable role in Cu homeostasis in plant. ...
Chapter
Heavy metals are required by plants in trace amounts for adequate growth and development, and their absence may lead to several detrimental effects during plant growth and development. Among them, one of the imperative trace elements required by plants for normal growth and development is copper. Copper (Cu) also serves as an important cofactor of many proteins. However, the details regarding these many Cu proteins are certainly limited. Nonetheless, the role played by these Cu proteins is of paramount significance. Cu holds an indispensable position in this regard; nevertheless, if its amount surpasses the required limit, it can lead to serious repercussions. Therefore, in order to maintain such a delicate balance, there exists an innate system within plants, which controls its absorption, distribution, and excretion within plants. There exists a unique set of proteins within plants termed as transport proteins, which regulate this delicate balance within plants. In the upcoming discussion, three of the most significant transport proteins also known as transporters are brought to light. These transport proteins include P type ATPases, that are responsible for the transport of Cu ions across the cell membrane, COPT proteins, that are responsible for the transport of Cu ions to various different cellular compartments, and chaperones that do not actually contain Cu but work like others, which are possessed with Cu. NRamp family gene analysis in soyabean seedlings also revealed their role during Cu and other heavy metal strain. The expression of this gene family also gets altered during heavy metal toxicity. The role of SPL7 transcription factor, in Cu homeostasis, has also been highlighted. In addition to it, the related role of Cu transport systems in biosynthesis and homeostasis has also been discussed.
... In response to BS11 inoculation, the copper (Cu) transporter protein 6 (spot GP3) was up-regulated in rice leaves compared to the control. Cu is an essential micronutrient for plants transported to various plant cells by copper transporter proteins (Puig 2014). It has been demonstrated in an earlier study that Cu is highly required for mitochondrial respiration, cell wall metabolism, photosynthesis, and oxidative stress protection in plants (Printz et al. 2016). ...
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In this study, we evaluated the efficacy of Bacillus strains for plant growth promotional activities under in vitro and in vivo conditions. The results indicated that Bacillus megaterium BS11 was superior in enhancing the plant growth and yield of rice plants compared to other Bacillus strains. Currently, there is no information available on the molecular mechanism of Rice–B. megaterium interaction for plant growth promotion. Thus, the present study was undertaken to understand the molecular basis of Rice–B. megaterium interaction at the proteome level using the two-dimensional polyacrylamide gel electrophoresis (2D-PAGE) approach. Proteomic results revealed that a total of 17 proteins were differentially expressed in response to BS11 inoculation. The application of BS11 up-regulated most of the identified proteins involved in plant metabolism, transcription, transporter, signaling, defense, and stress responses, which may underlie the improvement of rice plant growth. Furthermore, the proteomic results were validated at the mRNA level by Quantitative Real-Time PCR (qPCR) analysis. The abundances of proteins and transcripts were positively correlated for all genes except LOC_Os01g68620. Overall, our results suggest that B. megaterium strain BS11 may promote plant growth by improving various metabolism in rice plants.
... Helices 2, 5, and 6 are rich in aspartate residues and a signature domain is also characteristic for this family. The cytoplasm domain between helices 4 and 5 is rich in residual histidine and may be zinc-binding [77,83]. ...
... COPT are responsible for Cu 2+ ion collection and transport. Five COPT transporters (1-5) have been found in A. thaliana [65,83], with a higher shoot activity. COPT proteins occur in plasma membranes and transport copper from extracellular spaces to the cytoplasm or vacuole. ...
... COPT proteins occur in plasma membranes and transport copper from extracellular spaces to the cytoplasm or vacuole. The COPT protein level is regulated by copper ions [75,83]. ...
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The main mechanism of plant tolerance is the avoidance of metal uptake, whereas the main mechanism of hyperaccumulation is the uptake and neutralization of metals through specific plant processes. These include the formation of symbioses with rhizosphere microorganisms, the secretion of substances into the soil and metal immobilization, cell wall modification, changes in the expression of genes encoding heavy metal transporters, heavy metal ion chelation, and sequestration, and regenerative heat-shock protein production. The aim of this work was to review the natural plant mechanisms that contribute towards increased heavy metal accumulation and tolerance, as well as a review of the hyperaccumulator phytoremediation capacity. Phytoremediation is a strategy for purifying heavy-metal-contaminated soils using higher plants species as hyperaccumulators.
... Several studies have proven copper's diverse significance in biological systems. Copper's role in human disorders has been studied from a medicinal-chemical [17] and biochemical [18] standpoint, with a focus on the molecular physiology of Cu transport [19] copper homeostasis and its relationship to iron metabolism, as well as the role of copper in biological processes relevant to human physiology and illness, are the subjects of much contemporary research. While many of the activities claimed to explain inorganic noncomplexed copper homeostasis in humans have been outlined [19,20], only a few review studies have focused on the many biochemical events that could be directly implicated in the usage of copper complexes in medicine. ...
... Copper's role in human disorders has been studied from a medicinal-chemical [17] and biochemical [18] standpoint, with a focus on the molecular physiology of Cu transport [19] copper homeostasis and its relationship to iron metabolism, as well as the role of copper in biological processes relevant to human physiology and illness, are the subjects of much contemporary research. While many of the activities claimed to explain inorganic noncomplexed copper homeostasis in humans have been outlined [19,20], only a few review studies have focused on the many biochemical events that could be directly implicated in the usage of copper complexes in medicine. Some new bivalent metal complexes of Cu(II), Ni(II), and Co(II) with benzo thiazol ligands are examples of bioactive complexes. ...
Article
The physicochemical and antibacterial properties of compounds of copper and cobalt of the type (4PY-H)2[MCl4] and [M(Cl)2(4PY)2] were studied. The conventional coordination compounds and the acido complex crystals were prepared in a 1:2 mole ratio. The magnetic and electronic spectra data of the copper compounds showed that the two behave similarly to each other, even though, the copper acido complex had better absorptive properties. The copper acido complex crystalized in the monoclinic Ia space group, while the copper complex crystalized in the triclinic P1 space group. The cobalt complex similarly crystalized in the triclinic P1 space group while its acido complex crystalized also crystalized in a monoclinic crystal system, but contrary to the copper acido complex crystal, it was under P21/c space group. FTIR indicated that the compounds were structurally different where the acido complexes and the free ligand both displayed N-H peak in their IR spectra. This proved the existence of a protonated pyridyl nitrogen. Additionally, In silico geometry and FTIR data of all the 4 compounds corroborated well with the experimental data. The two compounds had very different melting points where the complex melted at higher temperatures than the acido complex. The compounds were thermally stable with melting points above 100 °C with complexes having higher melting points than acido complex crystals for both copper and cobalt. Also, DSC profiles showed that with increase in heat the acido complex crystals transformed to their complex forms. The antibacterial activity studies showed that the cobalt compounds were more active against the Gram-negative bacteria with more activity towards K. pneumoniae while the copper compounds were more active against the Gram-positive especially towards S. aureus. The time kill kinetics showed all the copper compounds to be bactericidal while the cobalt compounds were bacteriostatic.