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Search indexed bioRxiv preprints in genomics, neuroscience, cell biology and bioinformatics. Read source abstracts and check manuscript versions; preprints are not peer reviewed.

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At least 757 records · Page 42Linked to original sources

bioRxiv2020

Microfluidic platform for live cell imaging of 3D cultures with clone retrieval

Combining live imaging with the ability to retrieve individual cells of interest remains a technical challenge. These combined methods are of particular interest when studying highly dynamic or transient, asynchronous or heterogeneous cell biological and developmental processes. Here we present a method to encapsulate live cells in a 3D hydrogel matrix, via droplet compartmentalisation. Using a small-scale screen, we optimised matrix conditions for the culture and multilineage differentiation of mouse embryonic stem (ES) cells. Moreover, we designed a custom microfluidic platform that is compatible with live imaging. With this platform we are able to retain or extract individual bead/droplets by media flow only, obviating the need for enzymatic cell removal from the platform. We show that we can differentiate mES cells, monitor reporter expression by live imaging, and retrieve individual droplets for functional assays, correlating reporter expression with functional response. Overall, we present a highly flexible 3D cell encapsulation and microfluidic platform that enables both monitoring of cellular dynamics and retrieval for molecular and functional assays.

bioengineering↗

bioRxiv2020

I-KCKT allows dissection-free RNA profiling of adult Drosophila intestinal progenitor cells

The adult Drosophila intestinal epithelium is a model system for stem cell biology, but its utility is limited by current biochemical methods that lack cell type resolution. Here, we describe a new proximity-based profiling method that relies upon a GAL4 driver, termed intestinal-kickout-GAL4 (I-KCKT-GAL4), exclusively expressed in intestinal progenitor cells. This method used UV cross-linked whole animal frozen powder as its starting material to immunoprecipitate the RNA cargoes of transgenic epitope-tagged RNA binding proteins driven by I-KCKT-GAL4. When applied to the general mRNA-binder, poly(A)-binding protein, the RNA profile obtained by this method identified 98.8% of transcripts found after progenitor cell sorting, and had low background noise despite being derived from whole animal lysate. We also mapped the targets of the more selective RNA binder, Fragile Mental Retardation Protein, using enhanced CLIP, and report for the first time its binding motif in Drosophila cells. This method will therefore enable the RNA profiling of wildtype and mutant intestinal progenitor cells from intact flies exposed to normal and altered environments, as well as the identification of RNA-protein interactions critical for stem cell function. Summary StatementWe report a dissection-free method to identify proximity-based RNA-protein interactions in an in vivo stem cell population, enabling molecular analysis of these cells at unprecedented speed and resolution.

genetics↗

bioRxiv2020

Opposing effects of T cell receptor signal strength on CD4 T cells responding to acute versus chronic viral infection

A hallmark of the adaptive immune response is the ability of CD4 T cells to differentiate into a variety of pathogen appropriate and specialized effector subsets. A long-standing question in CD4 T cell biology is whether the strength of TCR signals can instruct one Th cell fate over another. The contribution of TCR signal strength to the development of Th1 and T follicular helper (Tfh) cells has been particularly difficult to resolve, with conflicting results reported in a variety of models. Although cumulative TCR signal strength can be modulated by the infection specific environment, whether or not TCR signal strength plays a dominant role in Th1 versus Tfh cell fate decisions across distinct infectious contexts is not known. Here we characterized the differentiation of CD4 TCR transgenic T cells responding to a panel of recombinant wild type or altered peptide ligand lymphocytic choriomeningitis viruses (LCMV) derived from acute and chronic parental strains. We found that while TCR signal strength positively regulates T cell expansion in both infection settings, it exerts opposite and hierarchical effects on the balance of Th1 and Tfh cells generated in response to acute versus persistent infection. The observation that weakly activated T cells, which comprise up to fifty percent of an endogenous CD4 T cell response, support the development of Th1 effectors highlights the possibility that they may resist functional inactivation during chronic infection. We anticipate that the panel of variant ligands and recombinant viruses described herein will be a valuable tool for immunologists investigating a wide range of CD4 T cell responses. Graphical abstract O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=192 SRC="FIGDIR/small/236497v2_ufig1.gif" ALT="Figure 1"> View larger version (50K): org.highwire.dtl.DTLVardef@1fdb592org.highwire.dtl.DTLVardef@19bc5e3org.highwire.dtl.DTLVardef@1dcf03forg.highwire.dtl.DTLVardef@20f27f_HPS_FORMAT_FIGEXP M_FIG C_FIG HighlightsO_LIIdentification of a wide panel of altered peptide ligands for the LCMV-derived GP61 peptide C_LIO_LIGeneration of LCMV variant strains to examine the impact of TCR signal strength on CD4 T cells responding during acute and chronic viral infection C_LIO_LIThe relationship between TCR signal strength and Th1 differentiation shifts according to the infection context: TCR signal strength correlates positively with Th1 generation during acute infection but negatively during chronic infection. C_LI

immunology↗

bioRxiv2022

Broad Recognition of Mamu-Bw4 and -A-related MHC class I Ligands by Rhesus Macaque Killer-Cell Immunoglobulin-Like Receptors

Definition of MHC class I ligands of rhesus macaque KIRs is fundamental to NK cell biology in this species as an animal model for infectious diseases, reproductive biology, and transplantation. To provide a more complete foundation for studying NK cell responses, rhesus macaque KIRs representing common allotypes of lineage II KIR genes were tested for interactions with MHC class I molecules representing diverse Mamu-A, -B, -E, -F, -I and -AG alleles. KIR-MHC class I interactions were identified by co-incubating reporter cell lines bearing chimeric KIR-CD3{zeta} receptors with target cells expressing individual MHC class I molecules and were corroborated by staining with KIR IgG-Fc fusion proteins. Ligands for 11 KIRs of previously unknown specificity were identified that fell into two general categories: interactions with multiple Mamu-Bw4 molecules or with Mamu-A-related molecules, including several allotypes of Mamu-AG and the hybrid Mamu-B*045:03 molecule. Although both groups include inhibitory and activating receptors, the majority of KIRs found to interact with Mamu-Bw4 are inhibitory, whereas most of the KIRs that interact with Mamu-AG are activating. We also identified Mamu-A1*012:01 as a ligand for KIR3DLw03*002, which belongs to a phylogenetically distinct group of macaque KIRs with a three amino acid deletion in D0 that is also present in human KIR3DL1/S1 and KIR3DL2. This study more than doubles the number of rhesus macaque KIRs with defined MHC class I ligands and identifies novel interactions with Mamu-AG, -B*045, and -A1*012. These findings support overlapping, but nonredundant, patterns of ligand recognition that reflects extensive functional diversification of these receptors.

immunology↗

bioRxiv2023

Anti-inflammatory role of APRIL by modulating regulatory B cells in antigen-induced arthritis

APRIL (A Proliferation-Inducing Ligand) acts as a secreted factor and plays important roles in B cell biology and chronic inflammation. The reported capacity of APRIL to stimulate B cells in vitro raised the hypothesis that APRIL may be a disease promoter in autoimmune diseases such as rheumatoid arthritis. In contrast, there is large evidence that B Lymphocyte Stimulator Factor (BAFF) can break B cell tolerance in mice and humans. This study was undertaken to examine the ability of APRIL to regulate B cell-mediated inflammatory response in a model of antigen-induced arthritis (AIA) in mice. AIA was induced by administration of antigen (mBSA) into the knee joints of previously immunized APRIL-transgenic (Tg) mice and their littermates. Flow cytometry was used to analyze the different inflammatory cell populations in spleen and draining lymph nodes in acute and chronic disease, while cytokine levels were assessed by ELISA. In the acute stage of disease APRIL-Tg mice displayed a decrease in joint swelling, disease score and inflammatory infiltrate in articular tissues, as compared with their littermates. In addition, we observed a decrease of cellularity in draining lymph nodes of APRIL-Tg mice. Flow cytometry analysis revealed an increase of CD19+IgM+CD5+ cell population in draining lymph nodes and increase of CD19+CD21hiCD23hi (B regulatory) cells in APRIL-Tg mice with arthritis and, increase of IL-10 and CXCL13 production in vitro.

immunology↗

bioRxiv2024

Artificial cells with all-aqueous droplet-in-droplet structures for spatially separated transcription and translation

The design of functional artificial cells involves compartmentalizing biochemical processes to mimic cellular organization. To emulate the complex chemical systems in biological cells, it is necessary to incorporate an increasing number of cellular functions into single compartments. Artificial organelles that spatially segregate reactions inside artificial cells will be beneficial in this context by rectifying biochemical pathways. In our study, we developed artificial cells featuring all-aqueous droplet-in-droplet structures that separate transcription and translation processes, mimicking the nucleus and cytosol in eukaryotic cells. This droplet-in-droplet architecture utilizes intrinsically disordered protein (IDP) to form coacervate droplets for the inner compartments, and aqueous two-phase systems (ATPS) for the outer compartments, with the outer interfaces stabilized by colloidal emulsifiers. The inner droplet was designed to enrich DNA and RNA polymerase to conduct transcription, which was coupled to translation at the outer droplet, realizing the cascade reaction mediated by mRNA. We also demonstrate that these processes proceed independently within each artificial cell compartment, maintaining the correspondence between genotype and phenotype. The modular configuration of these artificial organelles could be extended to other enzymatic reactions. Coupled with the ease of manufacturing these artificial cells, which only requires simple agitation in an all-aqueous mixture, this approach provides a practical and accessible tool for exploring complex systems of artificial organelles within large ensembles of artificial cells.

synthetic biology↗

bioRxiv2025

Refined cell transfer model reveals roles for Ascl2 and Cxcr3 in splenic localization of mouse NK cells during virus infection

Cell transfer experiments complement the rigorous investigation of antiviral and antitumor functions of natural killer (NK) cells. Success in these endeavors is enhanced by expansion of small numbers of input NK cells driven by viral antigens or homeostatic proliferation in immunodeficient hosts. In contrast, analysis of other NK-cell functions, including immunoregulation, are non-proliferative and require an intact immune system in recipient mice. We reveal poor persistence of conventional congenic (CD45.1) BoyJ NK cells following adoptive transfer in comparison to CRISPR-generated CD45.1+ (JAXBoy) NK cells. Reciprocal transfers between C57BL/6 and JAXBoy mice substantially improve seeding and maintenance of donor NK cells. Using this system, we confirm that CXCR3 re-positions NK cells in the white pulp of the spleen after infection, which is vital for immunoregulation. Moreover, we discovered that the transcription factor ASCL2 is required for recruitment of NK cells into the spleen and white pulp. These results provide improved tools and novel insights into NK cell biology. Key pointsO_LIJAXBoy are superior to conventional BoyJ mice for NK cell persistence after transfer. C_LIO_LICXCR3 repositions donor NK cells in T/B-zones of the spleen after infection. C_LIO_LIThe transcription factor ASCL2 is required for NK-cell recruitment to spleen white pulp. C_LI

immunology↗

bioRxiv2019

Long-term absence of Langerhans cells alters the gene expression profile of keratinocytes and dendritic epidermal T cells

Tissue-resident and infiltrating immune cells are continuously exposed to molecules derived from the niche cells that often come in form of secreted factors, such as cytokines. These factors are known to impact the immune cells biology. However, very little is known about whether the tissue resident immune cells in return also affect the local environment. In this study, with the help of RNA-sequencing, we show for the first time that long-term absence of epidermal resident Langerhans cells (LCs) led to significant gene expression changes in the local keratinocytes and resident dendritic epidermal T cells. Thus, immune cells might play an active role in maintaining tissue homeostasis, which should be taken in consideration at data interpretation.

immunology↗

bioRxiv2025

scPRINT-2: Towards the next-generation of cell foundation models and benchmarks

Cell biology has been booming with foundation models trained on large single-cell RNA-seq databases, but benchmarks and capabilities remain unclear. We propose an additive benchmark across a gymnasium of tasks to discover which features improve performance. From these findings, we present scPRINT-2, a single-cell Foundation Model pre-trained across 350 million cells and 16 organisms. Our contributions in pre-training tasks, tokenization, and losses made scPRINT-2 state-of-the-art in expression denoising, cell embedding, and cell type prediction. Furthermore, with our cell-level architecture, scPRINT-2 becomes generative, as demonstrated by our expression imputation and counterfactual reasoning results. Finally, thanks to our pre-training database, we uncover generalization to unseen modalities and organisms. These studies, together with improved abilities in gene embeddings and gene network inference, place scPRINT-2 as a next-generation cell foundation model.

bioinformatics↗

bioRxiv2026

The complete genome of the KOLF2.1J reference iPSC line

While induced pluripotent stem cells (iPSCs) have gained popularity in studying neurodegenerative diseases, the heterogeneity of stem cells used across studies impacts cross-study comparison. The iPSC Neurodegenerative Disease Initiative (iNDI) selected the KOLF2.1J cell line and prioritized its use as a reference standard for studying the effects of pathogenic variants on cell biology due to its stability and neutral neurodegenerative disease genetic risk. This cell line, and its derivatives expressing over 100 variants related to Alzheimers disease, Parkinsons disease, and other neurological diseases, are available for academic and industry access. Current genomic data analyses are limited by the use of a human reference genome that does not capture the complete genetic background of a given iPSC line. While in the future this issue may be partially mitigated by the creation of a comprehensive human pangenome, previous work has shown that generating custom genomes is of value both to characterize the variation present and to serve as a more appropriate genomic reference. Here, we generated and characterized a custom complete genome assembly from KOLF2.1J. Mapping of sequencing reads to a personalized diploid assembly results in more comprehensive mapping compared to traditional linear references (i.e GRCh38). In addition, we provide a comprehensive custom gene annotation along with isoform expression and differential methylation analyses across multiple cell types. The assembly and all additional data is browsable and publicly available. This resource will enable more accurate investigation of the KOLF2.1J cell line and any genomics data generated compared to using traditional generalized references, while also serving as a foundational approach for establishing custom reference assemblies for other high-value iPSC lines.

genomics↗

bioRxiv2026

RegDesign-Bench: Benchmarking computational methods for cell-type-specific regulatory DNA design

The de novo design of regulatory DNA sequences that drive cell-type-specific gene expression could enable precise gene control in synthetic biology, cell engineering and gene therapy. Various computational methods have been developed for this task, yet systematic comparisons that guide method selection for different design objectives remain limited. Here, we introduce RegDesign-Bench, a standardized framework for evaluating cell-type-specific cis-regulatory element design through a comprehensive set of computational analyses. Among the methods tested, strong predicted activity was not consistently accompanied by cell-type-selective accessibility or prominent cell-type-associated motif signals. Similarity to reference sequence distributions also did not ensure high predicted target-cell activity or specificity. These findings demonstrate that favourable performance on individual measurements does not necessarily extend across regulatory properties and library characteristics, revealing application-dependent strengths and limitations of current design methods. Overall, RegDesign-Bench offers a general-purpose computational framework for comparing and selecting methods to design regulatory DNA that drives cell-type-specific gene expression.

genomics↗

bioRxiv2018

Functional heterogeneity within the developing zebrafish epicardium

The epicardium is essential during cardiac development, homeostasis and repair and yet fundamental insights into its underlying cell biology, notably epicardium formation, lineage heterogeneity and functional cross-talk with other cell types in the heart, are currently lacking. In this study, we investigated epicardial heterogeneity and the functional diversity of discrete epicardial subpopulations in the developing zebrafish heart. Single-cell RNA-sequencing uncovered three epicardial subpopulations with specific genetic programmes and distinctive spatial distribution within the developing heart. Perturbation of unique gene signatures uncovered distinct functions associated with each subpopulation and established novel epicardial roles in cell adhesion, migration, and chemotaxis as a mechanism for recruitment of leukocytes into the heart. This work elucidates the mutual spatiotemporal relationships between different epicardial subpopulations and assigns unique function to each during cardiac development. Understanding which mechanisms cells employ to establish a functional epicardium and to communicate with other cardiovascular cell types during development will bring us closer to repairing cellular relationships that are disrupted during cardiovascular disease.

developmental biology↗

bioRxiv2018

The Allen Cell Structure Segmenter: a new open sourcetoolkit for segmenting 3D intracellular structures influorescence microscopy images

A continuing challenge in quantitative cell biology is the accurate and robust 3D segmentation of structures of interest from fluorescence microscopy images in an automated, reproducible, and widely accessible manner for subsequent interpretable data analysis. We describe the Allen Cell and Structure Segmenter (Segmenter), a Python-based open source toolkit developed for 3D segmentation of cells and intracellular structures in fluorescence microscope images. This toolkit brings together classic image segmentation and iterative deep learning workflows first to generate initial high-quality 3D intracellular structure segmentations and then to easily curate these results to generate the ground truths for building robust and accurate deep learning models. The toolkit takes advantage of the high-replicate 3D live cell image data collected at the Allen Institute for Cell Science of over 30 endogenous fluorescently tagged human induced pluripotent stem cell (hiPSC) lines. Each cell line represents a different intracellular structure with one or more distinct localization patterns within undifferentiated hiPS cells and hiPSC-derived cardiomyocytes. The Segmenter consists of two complementary elements, a classic image segmentation workflow with a restricted set of algorithms and parameters and an iterative deep learning segmentation workflow. We created a collection of 20 classic image segmentation workflows based on 20 distinct and representative intracellular structure localization patterns as a "lookup table" reference and starting point for users. The iterative deep learning workflow can take over when the classic segmentation workflow is insufficient. Two straightforward "human-in-the-loop" curation strategies convert a set of classic image segmentation workflow results into a set of 3D ground truth images for iterative model training without the need for manual painting in 3D. The deep learning model architectures used in this toolkit were designed and tested specifically for 3D fluorescence microscope images and implemented as readable scripts. The Segmenter thus leverages state of the art computer vision algorithms in an accessible way to facilitate their application by the experimental biology researcher. We include two useful applications to demonstrate how we used the classic image segmentation and iterative deep learning workflows to solve more challenging 3D segmentation tasks. First, we introduce the Training Assay approach, a new experimental-computational co-design concept to generate more biologically accurate segmentation ground truths. We combined the iterative deep learning workflow with three Training Assays to develop a robust, scalable cell and nuclear instance segmentation algorithm, which could achieve accurate target segmentation for over 98% of individual cells and over 80% of entire fields of view. Second, we demonstrate how to extend the lamin B1 segmentation model built from the iterative deep learning workflow to obtain more biologically accurate lamin B1 segmentation by utilizing multi-channel inputs and combining multiple ML models. The steps and workflows used to develop these algorithms are generalizable to other similar segmentation challenges. More information, including tutorials and code repositories, are available at allencell.org/segmenter.

cell biology↗

bioRxiv2024

Simultaneous and Dynamic Super-Resolution Imaging of Two Proteins in Arabidopsis thaliana using dual-color sptPALM

Super-resolution microscopy techniques have revolutionized cell biology by providing insights into single-molecule dynamics and nanoscale organization within living cells. However, the application of dynamic live-cell methods in plants remains limited by the availability of suitable fluorophores for simultaneous visualization of multiple proteins. To address this challenge, we implemented a dual-color single-particle tracking photoactivated localization microscopy (sptPALM) approach based on codon-optimized photoactivatable fluorescent proteins PA-GFP and PATagRFP. Recently, we demonstrated their individual performance in single-color experiments in Nicotiana benthamiana and Arabidopsis thaliana cells. Here, we establish their combined use for dual-color sptPALM, enabling the simultaneous tracking of two distinct protein species within the same plant cell. This approach provides a framework to investigate the coordinated dynamics, interactions, and spatial organization of multiple proteins in living plant cells.

plant biology↗

bioRxiv2025

Accidental and Regulated Cell Death in Yeast Colony Biofilms

The yeast species Saccharomyces cerevisiae is one of the most intensively studied organisms on the planet due to being an excellent eukaryotic model organism in molecular and cell biology. In this work, we investigate the growth and morphology of yeast colony biofilms, where proliferating yeast cells reside within a self-produced extracellular matrix. This research area has garnered significant scientific interest due to its applicability in the biological and biomedical sectors. A central feature of yeast colony biofilm expansion is cellular demise, which is onset by one of two independent mechanisms: either accidental cell death (ACD) or regulated cell death (RCD). In this article, we generalise a recently developed continuum model for the nutrient-limited growth of a yeast colony biofilm to include the effects of ACD and RCD. This new model involves a system of four coupled nonlinear reaction-diffusion equations for the yeast-cell density, the nutrient concentration, and two species of dead cells. Numerical solutions of the spatially one and two-dimensional governing equations reveal the impact that ACD and RCD have on expansion speed, morphology and cell distribution within the colony biofilm. Our results show good qualitative agreement with our own experiments.

microbiology↗

bioRxiv2017

Spatial modeling of the membrane-cytosolic interface in protein kinase signal transduction

The spatial architecture of signaling pathways and the inter-action with cell size and morphology are complex but little understood. With the advances of single cell imaging and single cell biology it becomes crucial to understand intracel-lular processes in time and space. Activation of cell surface receptors often triggers a signaling cascade including the activation of membrane-attached and cytosolic signaling components, which eventually transmit the signal to the cell nucleus. Signaling proteins can form steep gradients in the cytosol, which cause strong cell size dependence. We show that the kinetics at the membrane-cytosolic interface and the ratio of cell membrane area to the enclosed cytosolic volume change the behavior of signaling cascades significantly. We present a mathematical analysis of signal transduction in time and space by providing analytical solutions for different spatial arrangements of linear signaling cascades. These investigations are complemented by numerical simulations of non-linear cascades and asymmetric cell shapes.

systems biology↗

bioRxiv2021

Subcellular proteomics of dopamine neurons in the mouse brain reveals axonal enrichment of proteins encoded by Parkinson's disease-linked genes

Dopaminergic neurons modulate neural circuits and behaviors via dopamine release from expansive, long range axonal projections. The elaborate cytoarchitecture of these neurons is embedded within complex brain tissue, making it difficult to access the neuronal proteome using conventional methods. Here, we demonstrate APEX2 proximity labeling within genetically targeted neurons in the mouse brain, enabling subcellular proteomics with cell type-specificity. By combining APEX2 biotinylation with mass spectrometry, we mapped the somatodendritic and axonal proteomes of midbrain dopaminergic neurons. Our dataset reveals the proteomic architecture underlying proteostasis, axonal metabolism, and neurotransmission in these neurons. We find a significant enrichment of proteins encoded by Parkinsons disease-linked genes in striatal dopaminergic axons, including proteins with previously undescribed axonal localization. These proteomic datasets provide a resource for neuronal cell biology, and this approach can be readily adapted for study of other neural cell types.

neuroscience↗

bioRxiv2025

Mycobacterium tuberculosis effector protein PE5 hijacks the host CRL2 ubiquitin ligase complex

Mycobacterium tuberculosis (Mtb) is the leading infectious killer and infects one quarter of the global population. During infection, Mtb evade host immune responses via secreted effector proteins that interfere with, modulate, and protect from potent antibacterial responses. One such class of mycobacterial effector proteins are the PE/PPE proteins, a huge family of proteins encoded by an impressive 10% of the Mtb genome. Because some PE/PPEs have demonstrated roles in immune regulation and host cell interaction, and because of the sheer number of PE/PPEs in pathogenic mycobacteria ([~]170-200+ members depending on the species), they are thought to contribute to Mtb virulence and pathogenesis. However, the cellular functions of Mtbs 169 PE/PPE proteins have yet to be comprehensively characterized, at least in part because of their high GC content and large regions of extremely repetitive regions found both in DNA and amino acid sequences. One member of this family, PE5, is likely critical for mycobacterial pathogenesis, but its cellular and molecular functions, particularly within a host cell, are unexplored. We investigated the molecular functions of PE5 using affinity purification coupled mass spectrometry (AP-MS) and identified an interaction with the CRL2 complex, a host E3 ubiquitin ligase complex. PE5 interacts with CRL2 through its C-terminal Gly-Gly motif, which is bound by the CRL2 substrate receptor, KLHDC2. PE5 does not get ubiquitinated, but it is degraded upon being bound by KLHDC2. Interestingly, binding to PE5 increases the autoubiquitination of KLHDC2, but this autoubiquitination does not interfere with KLHDC2s ability to degrade known substrates or the ability of CRL2 to degrade substrates bound by other substrate receptors. Therefore, while PE5 binds to CRL2-KLHDC2 and does not get ubiquitinated itself, it does not appear to impede CRL2 activity. This study identifies a novel interaction between PE5 and a host ubiquitin ligase pathway, and it raises new questions about how PE5s interaction with the host modulates cell biology to promote Mtb virulence. Furthermore, it implicates CRL2 complexes in mediating cell-intrinsic host response to Mtb infection, a novel function for this ubiquitin ligase complex. Our results expand our understanding of how PE/PPEs may target innate immune responses and contribute to our knowledge of how uncharacterized PE/PPEs contribute to Mtbs virulence.

microbiology↗