Concurrent evaluation of autophagy induction and Burkholderia infection at the single cell level.
Abstract not provided.
Maes, Danae Maria↗
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Concurrent evaluation of autophagy induction and Burkholderia infection at the single cell level.
Abstract not provided.
Maes, Danae Maria↗
Targeting the Bet-Hedging Strategy with an Inhibitor of Bacterial Efflux Capacity Enhances Antibiotic Efficiency and Ameliorates Bacterial Persistence In Vitro
Persistence is a bet-hedging strategy in bacterial populations that increases antibiotic tolerance and leads to the establishment of latent infections. In this study, we demonstrated that a synthetic non-toxic taxane-based reversal agent (tRA), developed as an inhibitor of ABC transporter systems in mammalian cancer cells, enhanced antibiotic killing of persister populations from different pathogens, including Burkholderia, Pseudomonas, Francisella, and Yersinia. Acting as an inhibitor of bacterial efflux at 100 nM, tRA99020 enhanced antibiotic efficiency and suppressed the production of natural products of Burkholderia species polyketide synthase (PKS) function. We demonstrate that the metabolites produced by PKS in response to stress by different antibiotics act as inhibitors of mammalian histone deacetylase activity and stimulate cell death. Applying a single-molecule fluorescence in situ hybridization (smFISH) assay, we analyzed on a single-cell level the activation profiles of the persistence regulating pks gene in Burkholderia thailandensis treated with tRA99020 and antibiotics. We posit that a multi-pronged approach encompassing antibiotic therapies and inhibition of efflux systems and fatty acid catabolism will be required for efficient eradication of persistent bacterial populations.
59 BASIC BIOLOGICAL SCIENCES↗
Winter warming in Alaska accelerates lignin decomposition contributed by Proteobacteria
Background: In a warmer world, microbial decomposition of previously frozen organic carbon (C) is one of the most likely positive climate feedbacks of permafrost regions to the atmosphere. However, mechanistic understanding of microbial mediation on chemically recalcitrant C instability is limited; thus, it is crucial to identify and evaluate active decomposers of chemically recalcitrant C, which is essential for predicting C-cycle feedbacks and their relative strength of influence on climate change. Using stable isotope probing of the active layer of Arctic tundra soils after depleting soil labile C through a 975-day laboratory incubation, the identity of microbial decomposers of lignin and, their responses to warming were revealed. Results: The β - Proteobacteria genus Burkholderia accounted for 95.1% of total abundance of potential lignin decomposers. Consistently, Burkholderia isolated from our tundra soils could grow with lignin as the sole C source. A 2.2 °C increase of warming considerably increased total abundance and functional capacities of all potential lignin decomposers. In addition to Burkholderia , α - Proteobacteria capable of lignin decomposition (e.g. Bradyrhizobium and Methylobacterium genera) were stimulated by warming by 82-fold. Those community changes collectively doubled the priming effect, i.e., decomposition of existing C after fresh C input to soil. Consequently, warming aggravates soil C instability, as verified by microbially enabled climate-C modeling. Conclusions: Our findings are alarming, which demonstrate that accelerated C decomposition under warming conditions will make tundra soils a larger biospheric C source than anticipated.
54 ENVIRONMENTAL SCIENCES↗
Potential Biocontrol Activities of Populus Endophytes against Several Plant Pathogens Using Different Inhibitory Mechanisms
The plant microbiome can be used to bolster plant defense against abiotic and biotic stresses. Some strains of endophytes, the microorganisms within plants, can directly inhibit the growth of plant fungal pathogens. A previously isolated endophyte from wild Populus (poplar), WPB of the species Burkholderia vietnamiensis, had robust in vitro antifungal activity against pathogen strains that are highly virulent and of concern to Pacific Northwest agriculture: Rhizoctonia solani AG-8, Fusarium culmorum 70110023, and Gaemannomyces graminis var. tritici (Ggt) ARS-A1, as well as activity against the oomycete, Pythium ultimum 217. A direct screening method was developed for isolation of additional anti-fungal endophytes from wild poplar extracts. By challenging pathogens directly with dilute extracts, eleven isolates were found to be inhibitory to at least two plant pathogen strains and were therefore chosen for further characterization. Genomic analysis was conducted to determine if these endophyte strains harbored genes known to be involved in antimicrobial activities. The newly isolated Bacillus strains had gene clusters for production of bacillomycin, fengicyn, and bacillibactin, while the gene cluster for the synthesis of sessilin, viscosin and tolaasin were found in the Pseudomonas strains. The biosynthesis gene cluster for occidiofungin (ocf) was present in the Burkholderia vietnamiensis WPB genome, and an ocf deletion mutant lost inhibitory activity against 3 of the 4 pathogens. The new isolates lacked the gene cluster for occidiofungin implying they employ different modes of action. Other symbiotic traits including nitrogen fixation, phosphate solubilization, and the production of auxins and siderophores were investigated. Although it will be necessary to conduct in vivo tests of the candidates with pathogen-infected agricultural crops, the wild poplar tree microbiome may be a rich source of beneficial endophyte strains with potential for biocontrol applications against a variety of pathogens and utilizing varying modes of action.
59 BASIC BIOLOGICAL SCIENCES↗
Whole‐Cell MALDI‐ToF MS Coupled with Untargeted Metabolomics Facilitates Investigations of Microbial Chemical Interactions
Abstract The emergence of drug‐resistant pathogens necessitates the development of new countermeasures. In this regard, the introduction of probiotics to directly attack or competitively exclude pathogens presents a useful strategy. Application of this approach requires an understanding of how a probiotic and its target pathogen interact. A key means of probiotic‐pathogen interaction involves the production of small molecules called natural products (NPs). Here, we report the use of whole‐cell matrix‐assisted laser desorption/ionization time‐of‐flight (MALDI‐ToF) mass spectrometry to characterize NP production by candidate probiotics (mouse airway microbiome isolates) when co‐cultured with the respiratory pathogen Burkholderia . We found that a Bacillus velezensis strain inhibits growth of and elicits NP production by Burkholderia thailandensis . Dereplication of known NPs detected in the metabolome of this B. velezensis strain suggests that a previously unannotated bioactive compound is involved. Thus, we present the use of whole‐cell MALDI as a broadly applicable method for screening the NP composition of microbial co‐cultures; this can be combined with other ‐omics methods to characterize probiotic‐pathogen and other microbe‐microbe interactions.
Aiosa, Nicole↗
Interaction of Soil Microbes with Organoclays and their Impact on the Immobilization of Hg under Aerobic Conditions
Immobilization of mercury (Hg) leaching from bank soils of East Fork Poplar Creek (EFPC) is considered part of remediation strategies to mitigate the amount of Hg entering the creek. Different approaches are currently being evaluated, such as utilizing engineered sorbents to immobilize Hg species in EFPC bank soils. However, the influence of environmental microbes on the immobilization of Hg by sorbents is unknown. Organocation-modified phyllosilicate clay minerals (organoclays) are widely used as sorbents for the immobilization of contaminants. This study evaluates the interactions of Serratia marcescens and Burkholderia thailandensis with the sorbent Organoclay PM-199 and their impact on the immobilization of Hg under aerobic conditions. We evaluated the competitive binding of Hg between sorbents and selected microorganisms in a series of pure culture studies using bacterial strains identified in EFPC bank soil samples. Our results suggest that Hg sorption by Organoclay PM-199 is not significantly impacted by common soil bacteria present in EFPC, specifically Serratia marcescens and Burkholderia thailandensis, which are known to form biofilms. These findings suggest that sorbent amendments are an effective strategy for the remediation of Hg contamination in natural ecosystems.
54 ENVIRONMENTAL SCIENCES↗
Convergent reductive evolution and host adaptation in Mycoavidus bacterial endosymbionts of Mortierellaceae fungi
Intimate associations between fungi and intracellular bacterial endosymbionts are becoming increasingly well understood. Phylogenetic analyses demonstrate that bacterial endosymbionts of Mucoromycota fungi are related either to free-living Burkholderia or Mollicutes species. The so-called Burkholderia-related endosymbionts or BRE comprise Mycoavidus, Mycetohabitans and Candidatus Glomeribacter gigasporarum. These endosymbionts are marked by genome contraction thought to be associated with intracellular selection. However, the conclusions drawn thus far are based on a very small subset of endosymbiont genomes, and the mechanisms leading to genome streamlining are not well understood. The purpose of this study was to better understand how intracellular existence shapes Mycoavidus and BRE functionally at the genome level. To this end we generated and analyzed 14 novel draft genomes for Mycoavidus living within the hyphae of Mortierellomycotina fungi. We found that our novel Mycoavidus genomes were significantly reduced compared to free-living Burkholderiales relatives. Using a genome-scale phylogenetic approach including the novel and available existing genomes of Mycoavidus, we show that the genus is an assemblage composed of two independently derived lineages including three well supported clades of Mycoavidus. Using a comparative genomic approach, we shed light on the functional implications of genome reduction, documenting shared and unique gene loss patterns between the three Mycoavidus clades. We found that many endosymbiont isolates demonstrate patterns of vertical transmission and host-specificity, but others are present in phylogenetically disparate hosts. We discuss how reductive evolution and host specificity reflect convergent adaptation to the intrahyphal selective landscape, and commonalities of eukaryotic endosymbiont genome evolution.
59 BASIC BIOLOGICAL SCIENCES↗
Campylobacter jejuni uses energy taxis and a dehydrogenase enzyme for l -fucose chemotaxis
ABSTRACT Campylobacter jejuni is a leading cause of bacterial diarrhea worldwide, and infection in infants is associated with growth stunting in low- and middle-income countries. Approximately half of all C. jejuni isolates are asaccharolytic, while the remainder encode enzymes capable of l -fucose catabolism. Our previous study suggested that breastfed infants originating from sub-Saharan Africa and South Asia were colonized less frequently with l -fucose-metabolizing C. jejuni isolates compared to asaccharolytic strains. We also showed that FucX is an l -fucose dehydrogenase that binds to its NADP + cofactor even in the absence of l -fucose and that this enzyme is sufficient for l -fucose chemotaxis when introduced into strains lacking the metabolic pathway. This study indicates that FucX may influence l -fucose chemotaxis by increasing cellular NADPH/NADP + ratios, which subsequently affect the energy taxis components, CetABC that respond to shifting gradients of electron acceptors and donors. Furthermore, C. jejuni CetAB and CetAC can complement Escherichia coli aerotaxis. Our data support a model in which the shift in NADP + levels, impacted by the FucX dehydrogenase, affects chemotaxis in response to l -fucose. This swimming behavior can be phenocopied with a homologous l -fucose dehydrogenase from Burkholderia multivorans . Taken together, our work provides a possible explanation for why l -fucose-metabolizing C. jejuni isolates swim away from intestinal epithelial cells toward free fucose in the lumen where they are subsequently cleared by breastfed infants. IMPORTANCE In this study, we identify a separate role for the Campylobacter jejuni l -fucose dehydrogenase in l -fucose chemotaxis and demonstrate that this mechanism is not only limited to C. jejuni but is also present in Burkholderia multivorans . We now hypothesize that l -fucose energy taxis may contribute to the reduction of l -fucose-metabolizing strains of C. jejuni from the gastrointestinal tract of breastfed infants, selecting for isolates with increased colonization potential.
59 BASIC BIOLOGICAL SCIENCES↗
Highly Active Carbon–Platinum-Based Nanozymes: Synthesis, Characterization, and Immunoassay Application
Nanozymes (nanomaterials with intrinsic enzyme-like characteristics) have gained much attention for diagnostics and therapy due to their excellent enzyme-mimicking capability, great stability in environments, and facile and low-cost production. However, developing nanozymes with a high catalytic constant, K cat , has been challenging. Herein, we report a class of nanozyme-mimicking peroxidases, which are formed by depositing ultrasmall platinum nanoparticles (Pt NPs) 1–2 nm in size on the surface of hydrophilic nitrogen-doped carbon nanoparticles (CN NPs). These nanozymes defined as CN-Pt NPs show a high peroxidase-like activity with K cat values of 1.27 M·mL/s·g for 3,3,5,5′-tetramethylbenzidine (TMB) and 1.97 M·mL/s·g for hydrogen peroxide (H 2 O 2 ), respectively, which are at least one or two orders higher than many other reported carbon–noble metal-based nanozymes. Our developed CN-Pt NPs were further utilized in a colorimetric immunoassay as signal amplifiers for the biomarker detection of Burkholderia pseudomallei, a Gram-negative bacterial pathogen classified as a tier 1 select agent by the US CDC. In conclusion, the assay achieved lower limits of detection of 0.11 ng/mL in phosphate-buffered saline (PBS) and 0.16 ng/mL in human serum, when compared to many other assays in detecting the same biomarker.
37 INORGANIC, ORGANIC, PHYSICAL, AND ANALYTICAL CH↗
Dynamic nitrogen fixation in an aerobic endophyte of Populus
Biological nitrogen fixation by microbial diazotrophs can contribute significantly to nitrogen availability in non-nodulating plant species. In this study of molecular mechanisms and gene expression relating to biological nitrogen fixation, the aerobic nitrogen-fixing endophyte Burkholderia vietnamiensis, strain WPB, isolated from Populus trichocarpa served as a model for endophyte–poplar interactions. Nitrogen-fixing activity was observed to be dynamic on nitrogen-free medium with a subset of colonies growing to form robust, raised globular like structures. Secondary ion mass spectrometry (NanoSIMS) confirmed that N-fixation was uneven within the population. A fluorescent transcriptional reporter (GFP) revealed that the nitrogenase subunit nifH is not uniformly expressed across genetically identical colonies of WPB and that only ~11% of the population was actively expressing the nifH gene. Higher nifH gene expression was observed in clustered cells through monitoring individual bacterial cells using single-molecule fluorescence in situ hybridization. Through 15 N 2 enrichment, we identified key nitrogenous metabolites and proteins synthesized by WPB and employed targeted metabolomics in active and inactive populations. We cocultivated WPB Pnif-GFP with poplar within a RhizoChip, a synthetic soil habitat, which enabled direct imaging of microbial nifH expression within root epidermal cells. We observed that nifH expression is localized to the root elongation zone where the strain forms a unique physical interaction with the root cells. This work employed comprehensive experimentation to identify novel mechanisms regulating both biological nitrogen fixation and beneficial plant–endophyte interactions.
15 N-tracking metabolomics↗
Horizontal Gene Transfer to a Defensive Symbiont with a Reduced Genome in a Multipartite Beetle Microbiome
Symbiotic mutualisms of bacteria and animals are ubiquitous in nature, running a continuum from facultative to obligate from the perspectives of both partners. The loss of functions required for living independently but not within a host gives rise to reduced genomes in many symbionts. Although the phenomenon of genome reduction can be explained by existing evolutionary models, the initiation of the process is not well understood. Here, we describe the microbiome associated with the eggs of the beetle Lagria villosa, consisting of multiple bacterial symbionts related to Burkholderia gladioli, including a reduced-genome symbiont thought to be the exclusive producer of the defensive compound lagriamide. We show that the putative lagriamide-producing symbiont is the only member of the microbiome undergoing genome reduction and that it has already lost the majority of its primary metabolism and DNA repair pathways. The key step preceding genome reduction in the symbiont was likely the horizontal acquisition of the putative lagriamide lga biosynthetic gene cluster. Unexpectedly, we uncovered evidence of additional horizontal transfers to the symbiont’s genome while genome reduction was occurring and despite a current lack of genes needed for homologous recombination. These gene gains may have given the genome-reduced symbiont a selective advantage in the microbiome, especially given the maintenance of the large lga gene cluster despite ongoing genome reduction.
59 BASIC BIOLOGICAL SCIENCES↗
Bioactive exometabolites drive maintenance competition in simple bacterial communities
During prolonged resource limitation, bacterial cells can persist in metabolically active states of non-growth. These maintenance periods, such as those experienced in stationary phase, can include upregulation of secondary metabolism and release of exometabolites into the local environment. As resource limitation is common in many environmental microbial habitats, we hypothesized that neighboring bacterial populations employ exometabolites to compete or cooperate during maintenance and that these exometabolite-facilitated interactions can drive community outcomes. Here, we evaluated the consequences of exometabolite interactions over the stationary phase among three environmental strains: Burkholderia thailandensis E264, Chromobacterium subtsugae ATCC 31532, and Pseudomonas syringae pv. tomato DC3000. We assembled them into synthetic communities that only permitted chemical interactions. We compared the responses (transcripts) and outputs (exometabolites) of each member with and without neighbors. We found that transcriptional dynamics were changed with different neighbors and that some of these changes were coordinated between members. The dominant competitor B. thailandensis consistently upregulated biosynthetic gene clusters to produce bioactive exometabolites for both exploitative and interference competition. These results demonstrate that competition strategies during maintenance can contribute to community-level outcomes. It also suggests that the traditional concept of defining competitiveness by growth outcomes may be narrow and that maintenance competition could be an additional or alternative measure.
59 BASIC BIOLOGICAL SCIENCES↗
Bacterial diversity dynamics in microbial consortia selected for lignin utilization
Lignin is nature’s largest source of phenolic compounds. Its recalcitrance to enzymatic conversion is still a limiting step to increase the value of lignin. Although bacteria are able to degrade lignin in nature, most studies have focused on lignin degradation by fungi. To understand which bacteria are able to use lignin as the sole carbon source, natural selection over time was used to obtain enriched microbial consortia over a 12-week period. The source of microorganisms to establish these microbial consortia were commercial and backyard compost soils. Cultivation occurred at two different temperatures, 30°C and 37°C, in defined culture media containing either Kraft lignin or alkaline-extracted lignin as carbon source. iTag DNA sequencing of bacterial 16S rDNA gene was performed for each of the consortia at six timepoints (passages). The initial bacterial richness and diversity of backyard compost soil consortia was greater than that of commercial soil consortia, and both parameters decreased after the enrichment protocol, corroborating that selection was occurring. Bacterial consortia composition tended to stabilize from the fourth passage on. After the enrichment protocol, Firmicutes phylum bacteria were predominant when lignin extracted by alkaline method was used as a carbon source, whereas Proteobacteria were predominant when Kraft lignin was used. Bray-Curtis dissimilarity calculations at genus level, visualized using NMDS plots, showed that the type of lignin used as a carbon source contributed more to differentiate the bacterial consortia than the variable temperature. The main known bacterial genera selected to use lignin as a carbon source were Altererythrobacter , Aminobacter , Bacillus , Burkholderia , Lysinibacillus , Microvirga , Mycobacterium , Ochrobactrum , Paenibacillus , Pseudomonas , Pseudoxanthomonas , Rhizobiales and Sphingobium . These selected bacterial genera can be of particular interest for studying lignin degradation and utilization, as well as for lignin-related biotechnology applications.
59 BASIC BIOLOGICAL SCIENCES↗
A Comparative Metagenomic Analysis of Specified Microorganisms in Groundwater for Non-Sterilized Pharmaceutical Products
In pharmaceutical manufacturing, ensuring product safety involves the detection and identification of microorganisms with human pathogenic potential, including Burkholderia cepacia complex (BCC), Escherichia coli, Pseudomonas aeruginosa, Salmonella enterica, Staphylococcus aureus, Clostridium sporogenes, Candida albicans, and Mycoplasma spp., some of which may be missed or not identified by traditional culture-dependent methods. In this study, we employed a metagenomic approach to detect these taxa, avoiding the limitations of conventional cultivation methods. We assessed the groundwater microbiome’s taxonomic and functional features from samples collected at two locations in the spring and summer. All datasets comprised 436–557 genera with Proteobacteria, Bacteroidota, Firmicutes, Actinobacteria, and Cyanobacteria accounting for > 95% of microbial DNA sequences. The aforementioned species constituted less than 18.3% of relative abundance. Escherichia and Salmonella were mainly detected in Hot Springs, relative to Jefferson, while Clostridium and Pseudomonas were mainly found in Jefferson relative to Hot Springs. Multidrug resistance efflux pumps and BlaR1 family regulatory sensor-transducer disambiguation dominated in Hot Springs and in Jefferson. These initial results provide insight into the detection of specified microorganisms and could constitute a framework for the establishment of comprehensive metagenomic analysis for the microbiological evaluation of pharmaceutical-grade water and other non-sterile pharmaceutical products, ensuring public safety.
59 BASIC BIOLOGICAL SCIENCES↗
The Impact of Sorbent Amendments for Mercury Remediation on the Viability of Soil Microorganisms
Mercury (Hg) remediation across contaminated environments in the United States is an ongoing project. As part of the Hg cleanup strategy at East Fork Poplar Creek (EFPC), located in Oak Ridge, TN, the deployment of sorbents is considered. However, the impact of sorbents on soil microorganisms is poorly understood. In this study, we investigated the effect of sorbents on soil microorganism viability and biofilm formation to assess soil health during sorbent application for Hg remediation. We specifically investigated the effect of two engineered sorbents, Organoclay PM-199 and Organoclay MRM (which are manufactured from clay minerals formulated for various remediation applications), on two gram-negative organisms ( Serratia marcescens and Burkholderia thailandensis ) isolated from the Hg-contaminated EFPC bank soil. Pure cultures of S. marcescens or B. thailandensis were amended with 5% (w/v) and 25% (w/v) PM-199 and MRM, respectively, for 9 days. The samples were harvested, and bacterial cell viability was determined using a BacLight staining kit. Results showed that the growth of sorbent-amended S. marcescens was inhibited in contrast to that of unamended control. Furthermore, biochemical assays were used to analyze bacterial biofilm formation and integral biofilm components. Our results suggest that biofilm formation by sorbent-amended S. marcescens was negatively affected. In contrast, B. thailandensis amended with low concentrations of MRM showed enhanced growth and notable differences in biofilm morphology. These results suggest that the use of organoclay PM-199 and MRM at higher concentrations in field studies may hinder the growth of specific soil microorganisms.
54 ENVIRONMENTAL SCIENCES↗
Combatting melioidosis with chemical synthetic lethality
Burkholderia thailandensishas emerged as a nonpathogenic surrogate forBurkholderia pseudomallei, the causative agent of melioidosis, and an important Gram-negative model bacterium for studying the biosynthesis and regulation of secondary metabolism. We recently reported that subinhibitory concentrations of trimethoprim induce vast changes in both the primary and secondary metabolome ofB. thailandensis. In the current work, we show that the folate biosynthetic enzyme FolE2 is permissive under standard growth conditions but essential forB. thailandensisin the presence of subinhibitory doses of trimethoprim. Reasoning that FolE2 may serve as an attractive drug target, we screened for and identified ten inhibitors, including dehydrocostus lactone (DHL), parthenolide, and β-lapachone, all of which are innocuous individually but form a chemical-synthetic lethal combination with subinhibitory doses of trimethoprim. We show that DHL is a mechanism-based inhibitor of FolE2 and capture the structure of the covalently inhibited enzyme using X-ray crystallography. In vitro, the combination of subinhibitory trimethoprim and DHL is more potent than Bactrim, the current standard of care against melioidosis. Moreover, unlike Bactrim, this combination does not affect the growth of most commensal and beneficial gut bacteria tested, thereby providing a degree of specificity againstB. pseudomallei. Our work provides a path for identifying antimicrobial drug targets and for utilizing binary combinations of molecules that form a toxic cocktail based on metabolic idiosyncrasies of specific pathogens.
Science & Technology - Other Topics↗
Multilevel analysis between Physcomitrium patens and Mortierellaceae endophytes explores potential long‐standing interaction among land plants and fungi
SUMMARY The model moss species Physcomitrium patens has long been used for studying divergence of land plants spanning from bryophytes to angiosperms. In addition to its phylogenetic relationships, the limited number of differential tissues, and comparable morphology to the earliest embryophytes provide a system to represent basic plant architecture. Based on plant–fungal interactions today, it is hypothesized these kingdoms have a long‐standing relationship, predating plant terrestrialization. Mortierellaceae have origins diverging from other land fungi paralleling bryophyte divergence, are related to arbuscular mycorrhizal fungi but are free‐living, observed to interact with plants, and can be found in moss microbiomes globally. Due to their parallel origins, we assess here how two Mortierellaceae species, Linnemannia elongata and Benniella erionia , interact with P. patens in coculture. We also assess how Mollicute ‐related or Burkholderia ‐related endobacterial symbionts (MRE or BRE) of these fungi impact plant response. Coculture interactions are investigated through high‐throughput phenomics, microscopy, RNA‐sequencing, differential expression profiling, gene ontology enrichment, and comparisons among 99 other P. patens transcriptomic studies. Here we present new high‐throughput approaches for measuring P. patens growth, identify novel expression of over 800 genes that are not expressed on traditional agar media, identify subtle interactions between P. patens and Mortierellaceae, and observe changes to plant–fungal interactions dependent on whether MRE or BRE are present. Our study provides insights into how plants and fungal partners may have interacted based on their communications observed today as well as identifying L. elongata and B. erionia as modern fungal endophytes with P. patens .
59 BASIC BIOLOGICAL SCIENCES↗