Biology subjects

Zhu, C.

Publications and source records attributed to Zhu, C..

At least 19 recordsLinked to original sources

bioRxiv2026

Fine-tuning an intracellular allosteric pocket programs GPCR signaling bias

Biased signaling of G protein-coupled receptors (GPCRs) offers a route for precise therapeutics with fewer adverse effects. Intracellular allosteric ligands can impose such bias, yet how subtle chemical changes within these pockets reshape receptor-transducer coupling and downstream signaling remains unclear. Here, we systematically fine-tuned the intracellular allosteric pocket of neurotensin receptor 1 (NTSR1) through chemical modification of the {beta}-arrestin-biased ligand SBI-553. Minor changes to the ligand scaffold altered transducer-coupling preferences, generating distinct profiles of G-protein and {beta}-arrestin signaling. Cryo-electron microscopy structures of NTSR1-{beta}-arrestin2 complexes bound to designed allosteric probes revealed ligand-dependent remodeling of the intracellular cavity that reshapes the receptor-effector interface. Alanine scanning identified transducer-sensitive residues within and beyond the immediate ligand-contact region. In mice, SBI-553 and N065 produced distinct physiological profiles, with N065 improving survival in a lethal endotoxemia model. Finally, phosphoproteomic profiling revealed partially overlapping but distinct downstream signaling profiles among the three modulators. Together, these findings establish the intracellular allosteric pocket as a tunable determinant of transducer coupling and show that closely related modulators can generate distinct signaling and physiological profiles.

pharmacology and toxicology↗

bioRxiv2026

Increased EV yield and bioactivity of matrix-bound nanovesicles compared to conditioned culture medium-derived extracellular vesicles

Aim: Extracellular vesicles (EVs) are nanoparticles secreted by cells for cell-cell communication functions, such as wound repair, but low yields often limit their use. Several isolation methods have been explored to increase EV yield from conditioned culture medium (liquid-EVs); however, few studies compare isolation techniques and yield between liquid-EVs and matrix-bound nanovesicles (MBVs), a novel EV subtype embedded within the extracellular matrix. Methods: We isolate liquid-EVs and MBVs from mesenchymal stem cells (MSC) using size exclusion chromatography (SEC) and ultracentrifugation (UC) across various timepoints (14, 21, 24, 30 days). EVs are characterized by their yield, size, morphology, protein concentration, surface marker expression (CD9, CD81, ALIX), zeta potential, and influence on fibroblast wound closure. Results: We find that isolation method influences yield, where both SEC isolated liquid-EVs and MBVs yield higher particle counts and protein compared to UC. Interestingly, we find that regardless of isolation method, MBVs have a significantly higher yield than liquid-EVs from the same culture (15.6-fold higher using UC and 7.49-fold higher using SEC). Culturing for 24 days yielded the greatest number of MBV particles and protein. We performed a fibroblast scratch assay to assess bioactivity and found that liquid-EVs decreased wound closure compared to a medium control, whereas MBVs did not. Conclusion: Our work suggests SEC maximizes particle and protein yields for MSC-EVs. By achieving much higher yields of MBVs compared to liquid-EVs, and their promise as a wound repair strategy, we offer an alternative EV choice when EV yields are an issue towards translation.

bioengineering↗

bioRxiv2026

The Human Gut Cell Atlas v1.0: A Unified, Community-BuiltReference of Cellular Diversity Across the Gut

Single-cell RNA-sequencing (scRNA-seq) studies have transformed our understanding of cellular diversity in the human intestine, but differences in anatomical sampling, experimental processing and cell-type nomenclature limit comparison across datasets. Here, we present the Human Gut Cell Atlas (HGCA) V1, a community-curated reference of 944,390 healthy and disease-adjacent intestinal cells from 502 samples, 265 donors and 24 landmark studies, linked to detailed anatomical, clinical, and technical metadata. As this metadata is of critical importance to human biology, we developed MetaManager, an open source data management framework for standardized metadata collection and tracking. A five-level consensus taxonomy resolves 94 homeostatic cell identities and enables transferrable, high-resolution annotation of new gut datasets. By assembling cell identity, sampling metadata, and independent spatial validation, intestinal segment and radial tissue depth were revealed to systematically shape cellular composition and transcriptional state, enabling inference of the presence of sampling-dependent tissue niches, including lymphoid follicles. Reference mapping accurately and rapidly recovered cohort-level Crohn's disease and treatment signatures without de novo annotation, while mapping human intestinal organoids quantified cell-type-specific identity, regional specification, maturation, and transcriptional fidelity to native tissue. The HGCA V1 provides a standardized cellular, anatomical coordinate, and metadata system for reproducible, high-quality analysis of intestinal biology across healthy tissue, disease-associated remodelling, and experimental models, providing a continuously evolving community reference for cumulative discovery in human intestinal biology.

systems biology↗

bioRxiv2026

The impact of sex and ancestry on the molecular landscape of pancreatic adenocarcinoma

Background: Pancreatic ductal adenocarcinoma (PDAC) is projected to become the second leading cause of cancer-related mortality, with an increasing incidence and dismal 10% survival at 5 years despite best available care. PDAC is characterized by low tumour cellularity, recurrent driver mutation alterations, and substantial inter and intra-tumoural heterogeneity. The influence of individual genetic features such as sex and ancestry on tumour heterogeneity remains poorly understood. Methods: We assembled a cohort of PDAC across five datasets for a total of 7,527 tumours. Measures of genomic instability were assessed across sex and ancestry on a global level (SNV density and percent genome altered by copy number events), and at a gene level for recurrent events. KRAS variant frequencies, ploidy and mRNA abundance were also evaluated. Ancestry was assigned using ADMIXTURE from germline genotypes where available, and patient reported otherwise. Results: We observed ancestry-specific mutation patterns in global point mutation density, with higher SNV burden in African PDAC tumours and lower SNV burden in Asian PDAC tumours as compared to European tumours. We also identified an ancestry bias in the driver gene, GNAS, in which driver SNVs in GNAS was detected more frequently in those of Asian ancestry, and less frequently in those of African ancestry. No sex- or ancestry-associations were found for KRAS, but individual KRAS variants showed a trend towards ancestry-related variation. Conclusions: These findings demonstrate that ancestry contributes to the genomic heterogeneity of PDAC, shaping both global and gene-specific mutation patterns. This work underscores the importance of including diverse ancestries in PDAC molecular profiling efforts which may inform the development of tailored molecular diagnostics and targeted therapies.

genomics↗

bioRxiv2026

Antibodies targeting a conserved cryptic epitope at the influenza hemagglutinin head-stem interface via distinct binding modes

Influenza A virus continues to pose pandemic threats, as demonstrated by the recent global spread of highly pathogenic H5N1 strains. Consequently, the development and characterization of vaccines targeting zoonotic influenza subtypes have become a major focus of the field. In this study, we characterize two broadly reactive antibodies previously isolated from H5N1 vaccines. Although they have distinct immunoglobulin germline gene usage and binding modes, cryo-EM analysis shows that both antibodies share a highly conserved cryptic epitope at the interface of the hemagglutinin (HA) head and stem domains. The antigenicity of this epitope is influenced by natural amino acid variation at residue 293 of HA subunit 1, and its access is facilitated by acid-induced conformational change of HA. Nevertheless, both antibodies are non-neutralizing and possess only marginal protective efficacy against seasonal and H5N1 viruses in vivo, thus representing suboptimal vaccine responses. Overall, our findings expand the interactions of cross-reactive antibodies with an under-characterized HA epitope and provide insights for next-generation influenza vaccine design.

biochemistry↗

bioRxiv2026

Loss of TDP-43 function drives cryptic circular RNAs in neurodegenerative diseases

TAR DNA-binding protein 43 (TDP-43) is a key pathological hallmark of amyotrophic lateral sclerosis (ALS) and frontotemporal dementia (FTD) and a critical regulator of RNA splicing. While loss of TDP-43 induces aberrant splicing in linear transcripts, its impact on circular RNA (circRNA) biogenesis remains unexplored. Here, we show that TDP-43 depletion in human neurons induces widespread circRNA changes, especially upregulation of a distinct class of cryptic circRNAs that arise specifically upon loss of TDP-43. Some of these cryptic circRNAs incorporate cryptic exons derived from intronic sequences. Notably, these cryptic circRNAs exhibit greater stability than their corresponding linear RNA isoforms and accumulate progressively in neurons. Moreover, cryptic circRNAs are elevated in postmortem brain tissues from ALS, FTD and Alzheimer's disease (AD) patients. These findings reveal a previously unrecognized role for TDP-43 in repressing cryptic circRNA formation and establish these circRNAs as stable molecular signatures of TDP-43 dysfunction.

neuroscience↗

bioRxiv2026

A One-Step Chemoselective Strategy for Hydroxymethylcytosine Sequencing in DNA and RNA

5-Hydroxymethylcytosine (5hmC) is a relatively stable chemical mark formed by oxidation of 5-methylcytosine in DNA, while its RNA counterparts, 5-hydroxymethylcytidine (hm5C) and 2'-O-methyl-5-hydroxymethylcytidine (hm5Cm), have recently emerged as potential regulators of post-transcriptional processes. Existing 5hmC enrichment methods rely on anti-bodies or enzymatic {beta}-glucosyltransferase labeling, are largely DNA-specific, and are not readily applicable to RNA. Here we report bisulfite-assisted labeling by thiol sequencing (BALT-seq), a one-step chemoselective strategy that installs a biotin-thioether handle directly onto hydroxymethylcytosine in DNA or RNA under mild acidic bisulfite conditions, without using enzymes. Mechanistic studies show that thiol capture of the bisulfite-activated 5hmC intermediate competes with canonical cytosine-5-methylenesulfonate formation, yielding a stable adduct that DNA polymerases bypass efficiently and that resists APOBEC-mediated deamination. In mouse embryonic stem cells, BALT-seq produces genome-wide 5hmC profiles highly concordant with 5hmC-Seal, TAB-seq, and oxBS-seq, recovers expected gene-body and chromatin-state enrichment, and extends to low-input cell-free DNA. On RNA, BALT-seq enriches hm5C/hm5Cm-containing transcripts more effectively than antibody-based hMeRIP and reveals structured modification landscapes across tRNA species and LINE/LTR repeat families. BALT-seq thus establishes a unified, enzyme-free chemical platform for hydroxymethylcytosine profiling across DNA, cell-free DNA, and RNA.

genomics↗

bioRxiv2026

A positively selected microRNA controls a reversible aging program in striated muscle

Aging is the primary risk factor for most chronic diseases and is characterized in striated muscle by progressive functional decline, mitochondrial dysfunction, and chronic inflammation. The miR-128-1 locus resides within a positively selected haplotype on chromosome 2q21.3 associated with variation in grip strength, pulmonary function, and cardiometabolic traits in humans. Here, we show that antisense oligonucleotide-mediated inhibition of miR-128-3p restores muscle mass and function in aged mice, improves cardiac function while limiting adverse remodeling following myocardial infarction, and ameliorates skeletal and cardiac muscle pathology in mouse and pig models of Duchenne muscular dystrophy. Across these contexts, miR-128-3p inhibition induces a conserved transcriptional response characterized by activation of mitochondrial programs and suppression of inflammatory and fibrotic signaling, resembling the effects of established longevity interventions. These findings identify miR-128-3p as a regulator of a conserved aging-associated program and establish its inhibition as a strategy to restore tissue function across aging-related muscle pathologies.

molecular biology↗

bioRxiv2026

Genome-wide evolutionary shifts shape the emergence of dominant circulating clones in Mycobacterium abscessus

Mycobacterium abscessus exhibits extensive genomic diversity, yet independently emerged dominant circulating clones (DCCs) have achieved widespread distribution. However, the evolutionary changes associated with their emergence and subsequent diversification remain poorly understood. Here, we analyzed 11,314 genomes from 30 countries using a conservative framework integrating population-wide validation and read-supported sequence reconstruction, defining a stable core genome of 3,001 genes. Across seven DCCs, accessory-gene gains consistently exceeded losses on the ancestral branches leading to clone formation, with recurrent functions involving metal homeostasis, metabolism, environmental sensing and stress responses. Recombination also contributed substantial variation in specific lineages; in DCC3, ancestral recombinant regions encompassed core genes involved in iron acquisition, respiratory metabolism and protein homeostasis. Following DCC establishment, the relative contribution of homologous recombination to mutation consistently declined during within-clone diversification, indicating a broad shift toward mutation-dominated core-genome evolution. Together, these findings suggest a recurring pattern in DCC evolution, with accessory-genome gain and lineage-specific recombination occurring during clone formation, followed by increasingly mutation-dominated diversification after establishment.

genomics↗

bioRxiv2026

Functional evaluation of a natural AAV capsid liver targeting motif in human hepatocytes

Background & Aims: Adeno-associated virus (AAV) vectors are attractive delivery vehicles for therapeutic gene delivery, and a notable feature of most AAVs is their natural tropism for the liver, which leads to significant hepatic uptake following systemic administration. In previous work, we identified 266G as a conserved motif on a variable region on the capsid of many commonly used AAV variants that controls liver uptake in both mice and non-human primates. This single amino acid could be functionally leveraged to engineer AAVs to either de-target from or enhance tropism to the liver. Here, we explored whether these observations extended to the human context. Methods: Two human hepatocyte models were tested: Fah-/-/Rag2-/-/Il2rg-/- (FRG) mice with humanized livers and a bioengineered human microliver platform in vitro. A barcoded AAV capsid library including standard control serotypes were used to assess the role of the 266G motif on gene transfer and transgene expression in both liver systems. Results: In vivo, 266G containing AAVs indeed targeted human hepatocytes superiorly, with some noted dependency on the degree of human-hepatocyte replacement in the chimeric mouse model. Initial studies in the micropatterned primary human hepatocyte co-culture model however demonstrated enrichment of heparin-binding AAVs, and not 266G variants. Notably, incorporation of polyethylene glycol (PEG) into the system modified the AAV transduction potential of those capsids including the liver-targeting motif, recapitulating the hepatocyte transduction pattern observed in vivo. Importantly, when PEG was used, the two human models, both at the DNA and RNA level, did correlate significantly. Conclusions: Our results showed the potential of a combinatorial AAV library for model validation and revealed the human microliver platform-PEG as a reliable system for the development of AAV therapeutics.

molecular biology↗

bioRxiv2026

Resolving the immune response across clonal cancer evolution in situ with whole transcriptome profiling

Single-cell spatial transcriptomics is now central to studying tumors in their native tissue context. Here we present the first comprehensive, independent evaluation of Atera, a new spatial whole-transcriptome platform, compared with Xenium in human ductal carcinoma in situ (DCIS). We show that Atera enables granular cell-state annotation and resolves rare cell populations, experimentally validated by multiplex immunofluorescence (IF). We further show that its transcriptome-wide coverage enables inference of copy-number alterations at single-cell resolution, allowing us to reconstruct the clonal evolution of DCIS. We orthogonally confirm the inferred copy-number alterations by whole-genome sequencing of 16 microdissected tumor regions from consecutive tissue sections. Finally, by mapping the immune microenvironment onto this clonal architecture, we demonstrate the feasibility of tracking the changes in immune response along the clonal tumor evolution in situ. Together, our results establish Atera as a validated platform for tracking clonal evolution and immune adaptation in clinical samples.

bioinformatics↗

bioRxiv2026

Parallel Emergence and Adaptive Evolution of Multicountry Circulating Mycobacterium abscessus Clones

Mycobacterium abscessus (MAB) is a recently emerged bacterial pathogen causing an increasing number of human infections. Despite increasing recognition that different circulating clones contribute disproportionately to MAB population expansion, the global population dynamics and the evolutionary pressures underlying the emergence and spread of MAB clones remain incompletely understood. We analyzed 11,314 publicly available genomes of MAB isolates sampled from 30 countries and identified 38 recently emerged clones that have spread across multiple countries, including the seven previously described Dominant Circulating Clones (DCCs). There was a significant association between the time since the emergence of each circulating clone and its population size, with older clones generally having larger populations. Among the 26 circulating clones of M. abscessus subsp. abscessus, 12 carried the macrolide-susceptible erm(41) 28C genotype, highlighting clone-level variation in macrolide susceptibility. Genome-wide analyses identified 55 genes that were previously under purifying selection but showed evidence of positive selection during recent clade expansion. Collectively, our findings reveal that the recent global expansion of MAB comprises a broad continuum of newly emerged clones, accompanied by widespread shifts in gene-level selective pressures indicative of ongoing adaptation to new ecological environments.

microbiology↗

bioRxiv2026

Single-cell analysis of chromatin accessibility and gene expression in Drosophila melanogaster embryos following hypoxia treatment

Limited oxygen supply or hypoxia can impair fetal development and lead to developmental disorders, but the molecular mechanism underlying this phenomenon remains poorly understood. It is also well known that hypoxia results in transcriptomic alterations and epigenetic reprogramming. Drosophila melanogaster (fruit fly) has been used for decades as a powerful model to dissect the molecular mechanisms regulating development. To better understand the role of early hypoxic stress on development, we performed single-cell joint analysis of chromatin accessibility and transcriptome to characterize the influence of hypoxia on Drosophila embryonic development. We identified hypoxia-induced alterations in both gene expression and chromatin accessibility across 22 cell groups, especially in the genes regulating organogenesis and development of neuronal, tracheal, and muscular systems, including a reduction of germ cells, suggesting a long-lasting influence of hypoxic stress at an early embryonic stage on development and reproduction. In summary, this study demonstrates that early embryonic hypoxia induces cell type- and dose-dependent changes in chromatin accessibility and gene expression, leading to distinct developmental phenotypic responses, such as reduced number of germ cells under both 3% and 5% O2. We further conclude that the tramtrack (ttk) gene is critical in germ cell development and reproduction in Drosophila melanogaster.

developmental biology↗

bioRxiv2026

Multiscale harmonization and semantic integration of biomedical data enable biological insights through immersive exploration

Single-cell atlassing efforts like the Human BioMolecular Atlas Program (HuBMAP) and the Cellular Senescence Network (SenNet) are producing multiscale datasets across the healthy, adult, human body, but this data is typically explored only on 2D screens with limited 3D affordances, even though understanding a cells location within a tissue, organ, and body requires reasoning across many orders of spatial magnitude. The Human Reference Atlas (HRA) provides standard terminologies and a Common Coordinate Framework (CCF) for harmonizing such data spatially and semantically. Building on the HRA Organ Gallery in virtual reality (VR) application, we present "HRA: Powers of Ten," which integrates, harmonizes, and visualizes biological data from these efforts immersively in VR using a Multiscale Elevator System that lets users descend, like riding an elevator through an inverted skyscraper, from a whole body view of 81 reference organs to datasets across 5 organs (lymph node, brain, large intestine, small intestine, liver), 5 assay types (CODEX, Visium, v-CyCIF, Xenium, SBF-SEM), and 4 spatial scales. Contributed by Data Providers, these VR scenes enable biological insights into senescence patterns, cellular neighborhoods, 3D tissue reconstruction, and subcellular liver architecture. A standard operating procedure supports adding further datasets. Freely available on the Meta Store to over 20 million headset owners, the application, data, and code are open-source.

bioinformatics↗

bioRxiv2026

PTPRJ Drives Clonal Selection in CEBPA mutated AML

Transcription factor CEBPA is mutated in 10-15% of acute myeloid leukaemia (AML), a haematopoietic malignancy with high mortality. CEBPA mutations show a distinct pattern, and most patients are biallelic, carrying both an in-frame C-terminal mutation and a frameshift N-terminal mutation on opposing alleles. Rare N-terminal germline cases have 100% penetrance to AML, all with an acquired a C-terminal mutation. This suggests a selective pressure from one CEBPA mutation to develop another. Our zebrafish models faithfully recapitulate the human disease. All biallelic mutant combinations die by 4-6 weeks of age, with pre-leukaemic haematopoietic stem cell (HSC) expansion. C-terminal and N-terminal mutants show phenotypic differences in myeloid-primed HSC and differences in the myeloid differentiation block. RNA-Seq identified differentially expressed genes in opposing vectors. We identified phosphatase receptor PTPRJ as a candidate driver of clonal selection, with knock-out of ptprja in our fish accelerating pre-leukaemic expansion of HSC in C-terminal mutants, decelerating it in N-terminal mutants. Cebpa-mutant murine cells exhibit changes in differentiation and a clonal advantage with loss of Ptprj, which perturbs key signalling pathways. Our data suggest that PTPRJ contributes to the mechanism of leukaemogenesis in CEBPA mutant AML by driving the selective pressure from each mutation to develop the other.

cancer biology↗

bioRxiv2026

HnRNPAB-targeted antisense oligonucleotides ameliorate tau pathology and cognitive deficits by modulating Alzheimers disease-associated alternative splicing

Aberrant alternative splicing alters multiple disease-associated splicing events, including MAPT exon 10 inclusion linked to tau pathology, and thereby impacts Alzheimers disease (AD) progression. Heterogeneous nuclear ribonucleoprotein AB (hnRNPAB) functions as a splicing regulator, yet its roles in AD remain poorly defined. Here, we identified AB332, the full-length isoform of hnRNPAB, as a novel repressor of MAPT exon 10 inclusion. AB332 binds a conserved AAUAU motif and recruits RBMX and RBMXL1 through its RRM and glycine-rich domains to assemble a complex. Beyond MAPT, AB332 modulates a network of splicing events in multiple AD-associated genes, including STAG2, ApoER2, MCL-1, PICALM, and zDHHC7. Importantly, the exon 7-skipping isoform AB285 lacks these activities. Transcriptomic analysis of post-mortem brain tissues from AD patients revealed reduced expression of hnRNPAB, while selective downregulation of AB332 was confirmed in the hippocampus of 3xTg AD mice. We performed a systematic antisense oligonucleotide-tiling screen targeting hnRNPAB exon 7 and its flanking intronic regions, identifying ASO30 that specifically upregulated AB332 expression. In 3xTg AD mice, ASO30 restored hippocampal AB332 levels, reduced pathological tau phosphorylation, and rescued cognitive deficits. These findings define a molecular mechanism by which AB332 regulates AD-associated splicing events, thereby providing a therapeutic strategy targeting hnRNPAB for AD. GRAPHICAL ABSTRACT O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=79 SRC="FIGDIR/small/736008v1_ufig1.gif" ALT="Figure 1"> View larger version (23K): org.highwire.dtl.DTLVardef@621096org.highwire.dtl.DTLVardef@2f01b1org.highwire.dtl.DTLVardef@182e6forg.highwire.dtl.DTLVardef@190afd3_HPS_FORMAT_FIGEXP M_FIG C_FIG

molecular biology↗

bioRxiv2026

CryoGO enables high-resolution structural profiling of endogenous cellular macromolecules

Resolving macromolecular structures within their native cellular environment is essential for connecting molecular architecture to physiological function, yet the field lacks accessible, high-throughput methods for generating suitable specimens from cells. Here, we introduce cryoGO (on-Grid Opening cryo-electron microscopy), a simple, rapid, and scalable strategy that mechanically opens cells directly on EM grids to produce cell-derived specimens for high-resolution single-particle cryo-EM. This approach enables near-atomic structure determination of diverse endogenous macromolecular assemblies spanning a broad molecular-weight range. We show that cryoGO captures the rich ribosomal conformational and compositional landscape while preserving physiological state distributions and aspects of spatial heterogeneity originating from cells. Furthermore, this rapid workflow enables time-resolved structural profiling, capturing both long-term cellular adaptation and acute remodeling on timescales of seconds. Compatible with standard cryo-EM infrastructure and diverse biological samples, while requiring only small numbers of cells, cryoGO lowers the technical barrier to high-resolution native structural biology.

biophysics↗

bioRxiv2026

Visualizing immunoreceptor forces and their effects in vivo

Immunoreceptors experience forces that modulate their activities; however, demonstrating this in vivo has been limited by technical challenges. As a first step toward meeting this challenge, we adapted a synthetic Notch (SynNotch) receptor system to report force transmission through immunoreceptors in vivo by replacing the native ligand-binding domain with a receptor-specific antibody and rewiring Notch signaling to drive EGFP or luciferase expression. We expressed SynNotch on Jurkat cells targeting CD40 or T cell receptor (TCR) and characterized their activation in coculture with B or T cells, defining the requirements, optimal conditions, and kinetics of activation. Using complementary mechanobiology approaches, we quantified the exogenous force required for reporter activation and verified that activation depends on forces generated by receptor-expressing sender cells rather than SynNotch-expressing receiver cells. By implanting sensors and targets into immunocompromised mice, we visualized mechanically activated reporter expression on CD40 and TCR-targeting SynNotch cells in vivo. Furthermore, CD40 and TCR signaling was amplified when the receptor bore force against mechanical support from immobilized ligand, indicating that force functions as biologically relevant co-stimulus. Together, our results establish mechanically activated SynNotch reporters as a useful strategy for detecting receptor-associated mechanical signaling across 2D coculture, 3D organoid, and in vivo systems.

bioengineering↗