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Atmospheric nitrogen (N) deposition could affect the structure and function of terrestrial plants. Non-N2-fixing lichens are used to monitor atmospheric N deposition because they rely on the deposited inorganic N (i.e., ammonium and nitrate) as N sources. However, the uptake capacities of lichen on ammonium and nitrate remain unclear, which hinders the application of lichen N content to accurate bioindication of atmospheric N deposition levels. We investigated ammonium and nitrate uptake capacities of Cladonia rangiferina, which was treated with ammonium alone, nitrate alone, and ammonium and nitrate mixture solutions with different mixing ratios under light and dark conditions. The results showed that N uptake rates increased with ammonium and nitrate concentrations in solutions and generally followed the Michaelis-Menten saturation kinetics. Ammonium uptake of C. rangiferina showed higher values of affinity, and was more efficient than the nitrate uptake. Both rates and amounts of nitrate uptake decreased with increasing ratios of ammonium to nitrate in solutions, while ammonium uptake showed no substantial variations, indicating an inhibition of ammonium on nitrate uptake capability. The darkness significantly decreased the maximum uptake rate and efficiency of nitrate, but had much weaker effects on lichen ammonium uptake. These findings highlight the preference of lichen on ammonium as a key N uptake strategy. It is thus necessary to consider the main types of atmospheric inorganic N deposition when using lichens to monitor atmospheric N pollution levels and evaluate N deposition based on lichen ecophysiology.

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The endolichenic fungus Aspergillus sp. was isolated from the lichen Xanthoparmelia conspersa harvested in France. Aspergillus sp. was grown on a solid culture medium to ensure the large-scale production of the fungus with a sufficient mass of secondary metabolites. The molecular network analysis of extracts and subfractions enabled the annotation of 22 molecules, guiding the purification process. The EtOAc extract displayed an antiproliferative activity of 3.2 ± 0.4 µg/mL at 48 h against human colorectal cancer cells (HT-29) and no toxicity at 30 µg/mL against human triple-negative breast cancer (TNBC) cells (MDA-MB-231) and human embryonic kidney (HEK293) non-cancerous cells. Among the five prenylated compounds isolated, of which four are echinulin derivatives, compounds 1 and 2 showed the most important activity, with IC50 values of 1.73 µM and 8.8 µM, respectively, against HT-29 cells.

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Lichens are increasingly recognised as valuable bioindicators for environmental heavy metal pollution due to their sensitivity to spatial and temporal variations in pollution levels and their ability to adapt to diverse and often harsh habitats. This review initially examines the mechanisms of metal absorption in lichens, including particulate entrapment, ion exchange, and intracellular absorption, as well as their physiological responses to abiotic stressors such as heavy metal exposure and desiccation. In the latter part, we compile and synthesise evidence showing that secondary metabolites in lichens are significantly influenced by metal concentrations, with varying impacts across different species. Although extensive research has addressed the broader physiological effects of heavy metal hyperaccumulation in lichens, there remains a significant gap in understanding the direct or indirect influences of heavy metals on the lichen microbiome, possibly mediated by changes in secondary metabolite production. Our review integrates these aspects to propose new research directions aimed at elucidating the mechanisms underlying physiological responses such as resilience and adaptability in lichens. Overall, this review highlights the dynamic interplay between microbiome composition, secondary metabolite variation, and metal accumulation, suggesting that these factors collectively contribute to the physiological responses of lichens in polluted environments.

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Two novel strains, YIM 133132T and YIM 133296, were isolated from lichen samples collected from Yunnan Province, Southwest PR China. YIM 133132T and YIM 133296 are aerobic, Gram-staining-positive, non-motile actinomycetes. They are also catalase-positive and oxidase-negative, and YIM 133132T formed flat yellowish colonies that were relatively dry on YIM38 agar medium. Flat yellowish colonies of YIM 133296 were also observed on YIM38 agar medium. YIM 133132T grew at 25-35 °C (optimum 25-30 °C), pH 6.0-9.0 (optimum pH 7.0) and in the presence of 0-8% (w/v) NaCl. Phylogenetic analysis based on 16S rRNA gene sequences revealed that strains YIM 133132T and YIM 133296 represented members of the genus Luteipulveratus and exhibited high sequence similarity (96.93%) with Luteipulveratus halotolerans C296001T. The genomic DNA G+C content of both strains was 71.8%. The DNA-DNA hybridisation (dDDH) values between YIM 133132T and YIM 133296 were 85.1%, and the DNA-DNA hybridisation value between YIM 133132T and YIM 133296 and L. halotolerans C296001T was 23.4%. On the basis of the draft genome sequences, the average nucleotide identity (ANI) between strains YIM 133132T and YIM 133296 and L. halotolerans C296001T was 80.8%. The major menaquinones that were identified were MK-8(H4), MK-9 and MK-8(H2). The polar lipids were diphosphatidylglycerol and phosphatidylinositol. On the basis of the morphological, physiological, biochemical, genomic, phylogenetic and chemotaxonomic characteristics, strains YIM 133132T and YIM 133296 can be clearly distinguished from L. halotolerans C296001T, and the two strains represent a novel species for which the name L. flavus sp. nov. is proposed. The type strain is YIM 133132T (CGMCC= 1.61357T and KCTC= 49824T).

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Soil salinization is a major environmental threat to the entire terrestrial ecosystem. Lichens arose from the symbiosis of fungi and algae or cyanobacteria. They have a high tolerance to various extreme environments, including adaptation to saline-alkali habitats. Thus, lichens are pioneer species on saline-alkali soil. However, the separate resilience of the two symbiotic partners under saline-alkali conditions remains insufficiently understood. In this study, two representative symbiotic algae, Diplosphaera chodatii and Trebouxia jamesii, were studied for their physiological response to the saline-alkali stress by adjusting different concentrations of NaHCO3, together with their respective symbiotic fungi Endocarpon pusillum (terricolous lichen) and Umbilicaria muhlenbergii (saxicolous lichen). The results indicate that cell growth rate and biomass in all four cultures decreased in alkali-alkaline substrate, while cellular activities and ultrastructure were affected to a distinct extent. Compared with the symbiotic fungi, the algae were found to be more active in coordinating oxidative stress and lipid peroxidation damage under the saline-alkali stress. The antioxidant system of the alga was especially shown as a key adaptive trait and it provides an important strategy for species survival and persistence in arid saline-alkali desert. The specific survival ability of the lichen symbiosis relies on the stress resilience advantages of the symbiotic partners in combination. Our study provided new insights into understanding the adaptation of lichen symbiosis to desert saline-alkali soil, and the potential of lichen symbiotic algae in the future desert ecological restoration.

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Lichens are important components of high-latitude boreal and Arctic habitats. While stress tolerant, they are among the most sensitive ecosystem components to climate change, in particular, an increase in ultraviolet light (UV) arising from polar ozone depletion and deforestation. This study is the first to explore the effects of UV-B on gene expression in lichens to predict metabolic pathways involved in tolerance. Using transcriptome profiling and bioinformatic analyses, here we studied the effects of UV-B on gene expression in lichens using Lobaria pulmonaria (L.) Hoff. as a model species. UV-B exposure causes significant browning of the upper cortex of the thallus, which correlates to an increased expression of biosynthetic gene clusters involved in the synthesis of eu- and allomelanins and melanin precursors. Based on transcriptome analyses, we suggest that the biosynthesis of melanins and other secondary metabolites, such as naphthalene derivates, tropolones, anthraquinones, and xanthones, is a trade-off that lichens pay to protect essential metabolic processes such as photosynthesis and respiration. Expression profiles of general stress-associated genes, in particular, related to reactive oxygen species scavenging, protection of proteins, and DNA repair, clearly indicate that the mycobiont is the more UV-B-responsive and susceptible partner in lichen symbiosis. Our findings demonstrate that UV-B stress activates an intricate gene network involved in tolerance mechanisms of lichen symbionts. Knowledge obtained here may enable the prediction of likely effects on lichen biodiversity caused by climate change and pollution.

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Lichens have developed numerous adaptations to optimize their survival in various environmental conditions, largely by producing secondary compounds by the fungal partner. They often have antibiotic properties and are involved in protection against intensive UV radiation, pathogens, and herbivores. To contribute to the knowledge of the arsenal of secondary compounds in a crustose lichen species, we sequenced and assembled the genome of Toniniopsis dissimilis, an indicator of old-growth forests, using Oxford Nanopore Technologies (ONT, Oxford, UK) long reads. Our analyses focused on biosynthetic gene clusters (BGCs) and specifically on Type I Polyketide (T1PKS) genes involved in the biosynthesis of polyketides. We used the comparative genomic approach to compare the genome of T. dissimilis with six other members of the family Ramalinaceae and twenty additional lichen genomes from the database. With only six T1PKS genes, a comparatively low number of biosynthetic genes are present in the T. dissimilis genome; from those, two-thirds are putatively involved in melanin biosynthesis. The comparative analyses showed at least three potential pathways of melanin biosynthesis in T. dissimilis, namely via the formation of 1,3,6,8-tetrahydroxynaphthalene, naphthopyrone, or YWA1 putative precursors, which highlights its importance in T. dissimilis. In addition, we report the occurrence of genes encoding ribosomally synthesized and posttranslationally modified peptides (RiPPs) in lichens, with their highest number in T. dissimilis compared to other Ramalinaceae genomes. So far, no function has been assigned to RiPP-like proteins in lichens, which leaves potential for future research on this topic.

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Eighteen nitrogen-containing compounds (1-18) were isolated from cultures of the lichen-associated Streptomyces flavidovirens collected from the Qinghai-Tibet Plateau, including seven phenazine derivatives with three new ones, named subphenazines A-C (2-4), two new furan pyrrolidones (8-9), and nine known alkaloids. The structures were elucidated by spectroscopic data analysis, and absolute configurations were determined by single-crystal X-ray diffraction and ECD calculations. The phenazine-type derivatives, in particular compound 3, exhibited significantly better antineuroinflammatory activity than other isolated compounds (8-18). Compound 3 inhibited the release of proinflammatory cytokines including IL-6, TNF-α, and PGE2, and the nuclear translocation of NF-κB; it also reduced the oxidative stress and activated the Nrf2 signaling pathway in LPS-induced BV2 microglia cells. In vivo anti-inflammatory activity in zebrafish indicated that 3 inhibited LPS-stimulated ROS generation. These findings suggested that compound 3 might be a potent antineuroinflammatory agent through the regulation of the NF-κB/Nrf2 signaling pathways.

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The analysis of the interaction between main bionts (mycobiont and photobiont) in the lichen symbiosis delivers substantial information about their preferences in the selection of symbiotic partners, and their ecological preferences. The selectivity in the Parmelia genus has been defined as strong so far. However, data on this lichen genus, which includes several widely distributed species, are biogeographically limited. Therefore, using specialization indicators and extended sampling, in this study, we estimated the interactions between the main bionts of selected Parmelia spp., using two levels of estimation (species/OTU and haplotype). A comparison of mycobiont-photobiont interactions at different levels showed that considering only mycobiont species and Trebouxia OTUs, greater specialization is found, while Parmelia species studied in this work present a more generalistic strategy in photobiont choice when haplotypes are considered. Despite the uneven sampling of Parmelia species, the interpretation of specialization within species and individuals of the genus leads to a more precise and accurate interpretation of their adaptation strategies. Furthermore, the data from P. sulcata indicate the existence of a different pool of compatible haplotypes in some geographical regions compared to neighboring areas. This observation suggests the potential influence of climatic factors.

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Despite growing concerns regarding the long-range transport (LRT) and ecological risks of organophosphate esters (OPEs), information on the environmental behaviors of OPEs in polar terrestrial ecosystems remains inadequate. In the present study, 10 OPEs were analyzed in soil and vegetation samples collected from Fildes Peninsula, Antarctica. The OPE concentrations in Antarctic soils, mosses, and lichens ranged from 0.87 to 15.7 ng/g dry weight (dw), 9.8 to 113 ng/g dw, and 3.6 to 75.2 ng/g dw, respectively. Non-chlorinated OPEs predominated in terrestrial matrices, accounting for approximately 76 % of the OPE composition. Source identification indicated that OPE contamination in Antarctica likely resulted from local anthropogenic sources and LRT. Moreover, the bioaccumulation behavior of OPEs from soil to vegetation was assessed using bioconcentration factors (BCFs), revealing a significant non-linear trend of initial increase and subsequent decrease in BCFs relative to the lipophilicities of the octanol-air partition coefficient (log KOA) and octanol-water partition coefficient (log KOW). While low levels of OPEs in Antarctic terrestrial environments were reported in this study, their sustained inputs and potential ecological risks in polar regions warrant further attention.

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Accurate estimation of cryptogam biomass, encompassing bryophytes and lichens, is crucial for understanding their ecological significance. This estimation is conducted based on the strong correlations between mass and volume of cryptogams. However, mass-volume correlations vary among cryptogams because of their morphological differences. This problem can be solved using models that consider life forms that classify cryptogams based on morphological similarities. In this study, we investigated whether life form models improve cryptogam biomass estimation accuracy. The cryptogam mass-volume correlation of each life form was estimated using Bayesian linear models. The coefficients and intercepts of linear models differed between life forms, which was attributed to the morphological characteristics of each life form. Therefore, life form models can improve the accuracy of estimation models by incorporating morphological differences. However, taxonomic models that consider only the taxonomic difference (bryophytes vs lichens) demonstrated better overall estimation than the life form models, probably because of the ability of taxonomic models to capture systematic differences between bryophytes and lichens. Furthermore, these models may mitigate estimation errors related to morphological variations that cannot be adequately represented by life form types. Based on these results, we propose the appropriate use of estimation models.

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The formation of new ribosomes is tightly coordinated with cell growth and proliferation. In eukaryotes, the correct assembly of all ribosomal proteins and RNAs follows an intricate scheme of maturation and rearrangement steps across three cellular compartments: the nucleolus, nucleoplasm, and cytoplasm. We demonstrate that usnic acid, a lichen secondary metabolite, inhibits the maturation of the large ribosomal subunit in yeast. We combine biochemical characterization of pre-ribosomal particles with a quantitative single-particle cryo-EM approach to monitor changes in nucleolar particle populations upon drug treatment. Usnic acid rapidly blocks the transition from nucleolar state B to C of Nsa1-associated pre-ribosomes, depleting key maturation factors such as Dbp10 and hindering pre-rRNA processing. This primary nucleolar block rapidly rebounds on earlier stages of the pathway which highlights the regulatory linkages between different steps. In summary, we provide an in-depth characterization of the effect of usnic acid on ribosome biogenesis, which may have implications for its reported anti-cancer activities.

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Lichens are dual organisms, with one major mycobiont and one major photobiont in each lichen symbiosis, which can survive extreme environmental conditions in the Arctic. However, the diversity and distribution of lichen photobionts in the Arctic remain poorly understood compared to their mycobiont partners. This study explored the diversity of lichen mycobionts and photobionts in 197 lichen samples collected from the Ny-Ålesund region (Svalbard, High Arctic). The nuclear ribosomal internal transcribed spacer (ITS) regions were sequenced and phylogenetic analyses were performed. The relationships between mycobionts and photobionts, as well as the association patterns, were also investigated. A total of 48 species of lichen mycobionts (16 families, nine orders) and 31 species/lineages of photobionts were identified. These 31 photobiont species belonged to one class (Trebouxiophyceae) and five genera, including 22 species of Trebouxia, five species of Asterochloris, two species of Chloroidium, one species of Symbiochloris, and one species of Coccomyxa. The results indicated that most analyzed lichen mycobionts could associate with multiple photobiont species, and the photobionts also exhibited a similar pattern. The results provided an important reference dataset for characterizing the diversity of lichen mycobionts and photobionts in the High Arctic region.

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A lichen is a symbiotic association composed of a primary mycobionts and one or more photobionts living mutualistically together, forming a distinct morphological entity beneficial to their partnership and to other associated fungi, photobionts, and bacteria that collectively make up the lichen biome. The taxonomic identification of a lichen species often requires determination of the primary mycobiont's secondary metabolites, the key morphological characteristics of the thallus, and how it relates to other lichen species as seen in DNA phylogeny. This chapter covers lichens and their bionts, taxonomic identification, and their chemical constituents as exemplified by what is found in lichen biomes, especially those endemic to North America. Extraction and isolation, as well as updates on dereplication methods using mass spectrometric GNPS and NMR spectroscopic spin network fingerprint procedures, and marker-based techniques to identify lichens are discussed. The isolation and structure elucidation of secondary metabolites of an endolichenic Penicillium species that produces bioactive compounds will be described in detail.

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The presence or absence of lichens serves as an indicator of the condition of an ecosystem and the degree to which it is contaminated by various agents, such as agrochemicals and metals. Evaluating the use of lichens as bioindicators of agrochemical contamination could provide a more comprehensive perspective of current contamination levels. Monitoring was conducted over a 4-month period in two study areas: one was a well-conserved area contaminated by metals, and the other was an area surrounded by agricultural crops contaminated by agrochemicals. Data on the presence and abundance of lichens in each study area were recorded at 10 monitoring points, a procedure that was repeated 16 times (every 15 days), and concentrations of heavy metals and "organophosphate" agrochemicals in the lichens collected were measured by means of Inductively Coupled Plasma Atomic Emission Spectroscopy (ICP-OES) and Gas Chromatography (GC), respectively. Generalized linear mixed models were used to assess abundance and richness, while general linear mixed models were used to attain Shannon diversity and Simpson dominance indices. Moreover, a multivariate analysis was performed in order to compare the lichen communities in both areas. The results indicated differences between the area contaminated by metals and that contaminated by agrochemicals in terms of abundance and Simpson's dominance index, while no differences were found in the case of the richness and diversity models. The PERMANOVA analysis additionally showed differences between the lichen communities in the two areas. The results also demonstrated that Canoparmelia caroliniana bioaccumulated metals in both areas. The levels of barium, cadmium, and sodium were higher in the area contaminated by metals, while concentrations of chromium and copper were higher in the area contaminated by agrochemicals. Finally, the concentrations of agrochemicals were higher in the area contaminated by agrochemicals and included toxic substances such as Methylparathion and Parathion, which are prohibited in Ecuador. In conclusion, this research underscores the importance of lichens as precise indicators of environmental health and contamination by agrochemicals and metals.

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Melanin is a dark pigment from the group of phenolic or indole polymers with inherent biocompatibility and antioxidant capacity. In extremophilic lichen Lobaria pulmonaria, melanin is responsible for protective properties against hostile environments. Herein, the ability of melanin extracted from L. pulmonaria to counteract oxidative stress and related damages was studied in the mouse diaphragm, the main respiratory muscle. Initial in vitro experiments demonstrated ultraviolet (UV)-absorbing, antioxidant and metal chelating activities of melanin. This melanin can form nanoparticles and stabile colloidal system at concentration of 5 µg/ml. Pretreatment of the muscle with melanin (5 µg/ml) markedly reduced UV-induced increase in intracellular and extracellular reactive oxygen species (ROS) as well as antimycin A-mediated enhancement in mitochondrial ROS production accompanied by lipid peroxidation and membrane asymmetry loss. In addition, melanin attenuated suppression of neuromuscular transmission and alterations of contractile responses provoked by hydrogen peroxide. Thus, this study shed the light on the perspectives of the application of a lichen melanin as a protective component for treatment of skeletal muscle disorders, which are accompanied with an increased ROS production.

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Ore mineralizations in bedrock and their exploitation may have a negative impact on air quality of surrounding urban areas and, subsequently, on human health. This study uses lichens as bioindicators of atmospheric pollution to evaluate the spatial distribution of potentially toxic elements (PTEs) in the towns close to the massive sulfide deposits of the Iberian Pyrite Belt (IPB) in SW Spain. Altogether 89 native lichen samples of Xanthoria parietina were collected from the mining towns, control towns out of the reach of the mining activity, as well as from distal sampling sites. The samples were analyzed for 29 elements after acid digestion. The concentrations for Co, Ni, Cu, Zn, As, Rb, Mo, Cd, Sn, Sb, Cs, Ba, W, Tl, Pb, S, and Fe are significantly higher in the mining towns in comparison to the control towns. The ore mineral-associated PTEs, including Cu, Zn, As, Ba, and Pb, exhibit extreme concentrations in the urban areas close to the mining activity, and particularly in the small settlement of La Dehesa next to the mineral processing plant and the tailings pond. The distal samples confirm the decrease in the concentrations of all PTEs, and these samples present similar values as in the control areas. The results, point at increased bioaccumulation of PTEs in the lichen thalli of the adjacent urban areas, suggesting that the air quality of the adjacent urban areas is locally impacted by the massive polymetallic sulfide deposits which is enhanced by the mining activity. Therefore, monitoring the urban air quality is recommended.

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BACKGROUND: Lichens, traditionally considered as a simple partnership primarily between mycobiont and photobiont, are, in reality, complex holobionts comprised of a multitude of microorganisms. Lichen mycobiome represents fungal community residing within lichen thalli. While it is acknowledged that factors like the host lichen species and environmental conditions influence the structure of the lichen mycobiome, the existing research remains insufficient. To investigate which factor, host genus or location, has a greater impact on the lichen mycobiome, we conducted a comparative analysis of mycobiomes within Parmelia and Peltigera collected from both Turkey and South Korea, using high-throughput sequencing based on internal transcribed spacer region amplification. RESULTS: Overall, the lichen mycobiome was dominated by Capnodiales (Dothideomycetes), regardless of host or location. At the order level, the taxonomic composition was not significantly different according to lichen genus host or geographical distance. Hierarchical clustering of the top 100 abundant ASVs did not clearly indicate whether the lichen mycobiome was more influenced by host genus or location. Analyses of community similarity and partitioning variables revealed that the structure of the lichen mycobiome is more significantly influenced by location than by host genus. When analyzing the core mycobiome by host genus, the Peltigera mycobiome contained more ASV members than the Parmelia mycobiome. These two core mycobiomes also share common fungal strains, including basidiomycete yeast. Additionally, we used chi-squared tests to identify host genus-specialists and location-specialists. CONCLUSIONS: By comparing lichen mycobiomes of the same genera across different countries, our study advances our comprehension of these microbial communities. Our study elucidates that, although host species play a contributory role, geographic distance exerts a more pronounced impact on the structure of lichen mycobiome. We have made foundational contributions to understanding the lichen mycobiome occupying ecologically crucial niches. We anticipate that broader global-scale investigations into the fungal community structures will provide more detailed insights into fungal residents within lichens.

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Lichens have great potential as food, functional food additives or flavourings. The presence of specific substances with multiple biological activities is one of the characteristics of lichens. However, research on lichens as a food source or functional food additive is limited. The present study simulated, for the first time, the potential bioaccessibility of active compounds from 6 lichen species in simulated gastric and intestinal conditions. An in vitro digestion showed that the lichen substances had different bioaccessibility and stability during digestion. It was found that the application of some metabolic modulators significantly altered the accumulation of metabolites in most species. In addition, the study demonstrated the antimicrobial activity of the tested extracts as well as of 14 isolated lichen metabolites. These multi-directional studies demonstrate the potential of lichens in terms of their use as antimicrobial functional food additives.

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