bims-instec Biomed News
on Intestinal stem cells and chemoresistance in colon cancer and intestinal regeneration
Issue of 2026–05–03
nineteen papers selected by
Maria-Virginia Giolito, Université Catholique de Louvain



  1. Lipids Health Dis. 2026 Apr 30.
      Dysregulated lipid metabolism has emerged as a defining hallmark of colorectal cancer (CRC) progression, particularly in metastatic disease, where metabolic adaptation and immune evasion are tightly interconnected, as demonstrated in both murine models and human studies. Increasing evidence has demonstrated that alterations in lipid synthesis, uptake, transport, and oxidation not only sustain tumor bioenergetics but also actively remodel the tumor immune microenvironment. Key lipid metabolic regulators-including FASN, SREBP signaling, CD36-mediated lipid uptake, cholesterol metabolism, and fatty acid oxidation-coordinate oncogenic signaling and promote immunosuppressive states characterized by T-cell exhaustion, macrophage polarization, and ferroptosis resistance, on the basis largely of correlative and preclinical evidence. Recent advances in multiomics technologies, including single-cell and spatial transcriptomics, metabolomics, and lipidomics, have enabled high-resolution mapping of lipid-dependent immune niches within metastatic CRC (mCRC) lesions. These approaches reveal lipid metabolism as a central organizer of tumor-immune interactions and identify previously unrecognized metabolic vulnerabilities. In this review, we integrate current knowledge on lipid metabolic reprogramming in CRC with emerging multiomics insights, highlighting the mechanisms linking lipid metabolism, ferroptosis, gut microbiota interactions, and immune remodeling. We further discuss therapeutic strategies targeting lipid metabolic pathways and their potential synergy with immunotherapy. Collectively, the results of this work suggest that understanding lipid metabolism is a unifying framework for understanding mCRC biology and developing metabolism-guided therapeutic interventions.
    Keywords:  Colorectal cancer; Lipid metabolism; Tumor immune microenvironment
    DOI:  https://doi.org/10.1186/s12944-026-02956-9
  2. Methods Cell Biol. 2026 ;pii: S0091-679X(26)00081-6. [Epub ahead of print]206 81-93
      Ferroptosis is a regulated form of cell death driven by iron-dependent lipid peroxidation, with emerging relevance in cancer biology, particularly in therapy-resistant tumors such as colorectal cancer. Accurate and reproducible methods to monitor ferroptosis are essential for understanding its mechanisms and therapeutic potential. In this methodological paper, we present an optimized protocol for the evaluation of lipid peroxidation as a surrogate marker of ferroptosis in colon cancer cell models. Specifically, we describe an approach for quantifying malonylaldehyde and hydroxyalkenals (MDA+ 4-HDA), as key byproducts of lipid oxidation, using a colorimetric assay adapted for cell culture. This methodological framework provides a reliable basis for dissecting ferroptotic responses and for evaluating the activity of ferroptosis-inducing compounds in colorectal cancer research.
    Keywords:  4-HAs; Intestine; Lipids; MDA; Oxidation; Redox
    DOI:  https://doi.org/10.1016/bs.mcb.2026.02.013
  3. Biosensors (Basel). 2026 Mar 25. pii: 189. [Epub ahead of print]16(4):
      In colorectal cancer (CRC), cancer-associated fibroblasts (CAFs) and the fibrotic stroma generate form a dense stromal barrier that restricts the intratumoural exposure and spatial distribution of oxaliplatin. To enable local stromal remodelling of this pathological stromal compartment, we selected fibroblast activation protein (FAP) as a stromal target and co-assembled two amphiphilic conjugates, oncoFAP and retinoic acid (RA), into an FAP-directed RA nanoformulation termed LRAFAP. LRAFAP exhibited a uniform size distribution (107.1 ± 5.8 nm), remained stable for at least 7 d at 37 °C in PBS or serum-containing PBS, and showed accelerated esterase-responsive release. In a TGF-β-induced CAF-like model, LRAFAP markedly suppressed the expression of CAF activation-associated markers, reducing Fap and Acta2 mRNA levels by approximately 70% and 60%, respectively. In vivo, LRAFAP showed enhanced accumulation in CAF-enriched tumours and an increase in intratumoural oxaliplatin levels of approximately 2.5-fold relative to oxaliplatin alone. LRAFAP also reduced collagen deposition and CAF activation markers, and enhanced the antitumour efficacy of oxaliplatin while maintaining good tolerability. Collectively, these findings indicate that LRAFAP promotes local stromal remodelling and improves intratumoural oxaliplatin exposure, thereby enhancing the efficacy of oxaliplatin-based chemotherapy in CRC.
    Keywords:  cancer-associated fibroblast; colorectal cancer; extracellular matrix; fibroblast activation protein; retinoic acid
    DOI:  https://doi.org/10.3390/bios16040189
  4. Cell Stem Cell. 2026 Apr 30. pii: S1934-5909(26)00146-3. [Epub ahead of print]
      eIF4G2 (DAP5/NAT1) is a non-canonical translation initiation factor, but its role in homeostasis is unclear. Using inducible Eif4g2 knockout mice and intestinal organoids, we show that eIF4G2 loss collapses Lgr5+ intestinal stem cell (ISC) and secretory maturation programs while preserving villus architecture. Transcriptomic and single-nucleus multiome analyses reveal a durable fetal-like/regenerative state with YAP-TEAD activation and regenerative absorptive cells. Ribosome profiling identifies selective translation-efficiency loss among chromatin regulators, especially the KAT3 coactivators CREBBP and EP300, resulting in reduced KAT3 abundance and global histone acetylation; chemical KAT3 inhibition phenocopies this state. CUT&Tag and assay for transposase-accessible chromatin sequencing (ATAC-seq) demonstrate that reduced eIF4G2-KAT3 output drives locus-selective enhancer remodeling, with loss of adult ISC/Wnt-Notch elements and activation of TEAD-enriched fetal loci, without inflammatory or integrated stress response programs driving the transition. Fetal intestinal spheroids remain viable despite similar biochemical defects, highlighting a stage-specific requirement for translational buffering in maintaining adult identity.
    Keywords:  differentiation; eIF4G2 (NAT1/p97/DAP5); epigenetic gene regulation; histone modification; intestinal stem cell; translation initiation
    DOI:  https://doi.org/10.1016/j.stem.2026.04.006
  5. Dig Dis Sci. 2026 Apr 27.
       BACKGROUND: Colorectal cancer (CRC) exhibits extensive cellular heterogeneity and complex tumor microenvironment (TME) interactions, which influence tumor progression and treatment response. However, the precise interplay between epithelial cells and cancer-associated fibroblasts and its prognostic relevance remain incompletely understood.
    METHODS: We performed single-cell RNA sequencing on CRC and matched normal tissues, integrating the data with spatial transcriptomics. Cellular heterogeneity, developmental trajectories, and intercellular communication were analyzed to identify key epithelial and fibroblast subpopulations.
    RESULTS: Nine epithelial and nine fibroblast subpopulations were identified, with FGGY + epithelial cells and COL11A1 + fibroblasts markedly enriched in tumors. Intercellular communication analysis revealed a tumor-specific epithelial-CAF axis mediated by PPIA-BSG signaling, accompanied by strong spatial co-localization. Trajectory and copy number variation (CNV) analyses indicated progenitor-like PLK1 + epithelial cells contribute to tumor plasticity. The FGCS score, derived from six genes from these two subsets, stratified patients by stage and survival, acting as an independent prognostic factor.
    CONCLUSION: This study provides a comprehensive, high-resolution view of CRC cellular and microenvironmental organization, revealing how epithelial-fibroblast interactions shape tumor progression and spatial niches, offering insights into CRC biology and potential therapeutic targets.
    Keywords:  Colorectal cancer; Epithelial cell; Fibroblast; Single-cell RNA sequencing; Spatial transcriptomics
    DOI:  https://doi.org/10.1007/s10620-026-09923-5
  6. Neoplasia. 2026 Apr 30. pii: S1476-5586(26)00044-8. [Epub ahead of print]77 101314
      Chemoresistance remains a major cause of treatment failure in colorectal cancer (CRC), yet the metabolic mechanisms sustaining efflux-mediated drug resistance are not fully defined. Here, we identify ATP-citrate lyase (ACLY) as a metabolic regulator linking citrate-dependent acetyl-CoA production to epigenetic control of MDR1/ABCB1 expression. Using genetic and pharmacologic approaches, we show that ACLY catalytic activity contributes to the maintenance of histone acetylation at H3K9 and H4K16 and supports MDR1 transcription in CRC cells. Consistently, ACLY overexpression enhances, whereas its inhibition reduces, MDR1 expression and associated resistance-related transcriptional programs. In human CRC specimens, ACLY and MDR1 levels positively correlate, with a stronger association observed in advanced-stage tumors, supporting clinical relevance of this metabolic-epigenetic axis. Metabolic tracing with 13C-glucose suggests that perturbation of citrate flux influences ACLY-associated pathways and acetyl-CoA availability. In this context, vitamin C treatment reduces citrate-derived acetyl-CoA and ACLY phosphorylation and is associated with global histone deacetylation and decreased MDR1 expression in vitro and in KRAS-mutant patient-derived xenografts. Together, these findings highlight ACLY-dependent acetyl-CoA production as a potential metabolic vulnerability linked to epigenetic regulation of drug efflux programs in CRC. Targeting this metabolic-chromatin axis may represent a strategy to modulate MDR1-associated chemoresistance.
    Keywords:  ACLY; Ascorbate; Cancer; Chemoresistance; Epigenetic; KRAS; MDR-1; Metabolism
    DOI:  https://doi.org/10.1016/j.neo.2026.101314
  7. Proc Natl Acad Sci U S A. 2026 May 05. 123(18): e2519981123
      Inflammatory injury to the intestine triggers a reprogramming of the intestinal epithelium to a fetal-like state that drives rapid restoration of the epithelial barrier. Although the intestinal microbiota is a key modulator of inflammation, its role in influencing epithelial fetal-like stem cell reprogramming and consequent restitution remains unclear. Using irradiation (IR) injury as a model for small intestinal epithelium injury and repair, we found that the intestinal microbiota accelerated epithelial restitution by amplifying a repair-associated inflammatory response that promoted the emergence of fetal-like intestinal epithelial cells (IECs), marked by Ly6a and Clu. NOD2, the strongest genetic link to the development of Crohn's disease, was found to be expressed in fetal-like IECs following injury. Employing an ileal organoid model, we demonstrated that NOD2 activation by its peptidoglycan ligand potentiated an inflammatory gene signature characterized by interferon signaling, concurrent with enterocyte recovery. NOD2 deficiency exacerbated epithelial apoptosis following IR injury, whereas epithelial-specific NOD2 signaling promoted fetal-like IEC emergence and increased epithelial proliferation. Collectively, these findings reveal a pivotal role for the microbiota and NOD2-mediated microbial sensing in regulating fetal-like IEC fate after injury, thus contributing to the protective function of this microbial sensor during intestinal inflammation.
    Keywords:  NOD2; fetal-like reversion; inflammation; intestinal regeneration; microbiota
    DOI:  https://doi.org/10.1073/pnas.2519981123
  8. Nat Commun. 2026 Apr 25.
      Microsatellite-stable (MSS) colorectal cancers (CRC) are largely unresponsive to immune checkpoint inhibition (ICI). The MAYA trial used temozolomide (TMZ) in MGMT-silenced MSS mCRC, hypothesizing that TMZ-induced hypermutation could sensitize tumors to ICI; the primary endpoint was met, showing durable responses with TMZ plus ipilimumab and nivolumab. We perform integrated spatial, transcriptomic, and immune profiling of longitudinal tumor and blood samples from patients treated on the MAYA trial. Post-TMZ increases in tumor mutational burden associate with improved progression-free survival. Spatial profiling demonstrates that clinical benefit is greatest in permissive tumor microenvironments. Responders exhibit enrichment of cytotoxic T cells across tumor and stromal compartments, whereas non-responders display heterogeneous cellular neighborhoods, with fibroblasts in close spatial proximity to T cells, consistent with barriers to immune-mediated clearance. Longitudinal peripheral immune profiling shows that early upregulation of TIGIT and PD-1 following TMZ exposure predicts resistance. Together, these findings indicate that both mutational evolution and spatial immune architecture contribute to immune sensitization in MGMT-silenced MSS CRC. Clinical Trial Identification: NCT03832621.
    DOI:  https://doi.org/10.1038/s41467-026-72204-2
  9. Nat Metab. 2026 Apr 27.
      Squamous cell carcinomas (SCCs) originate in epithelial tissues of older individuals who have been exposed to environmental carcinogens. Despite overlapping clinical hallmarks, SCCs from different anatomic sites have different prognoses. Here we show that fibroblasts confer site-specific cues that determine SCC proliferation and invasion. Oral and lung fibroblasts have distinct lipid metabolism, transferring unique lipids to SCC cells that promote epithelial-to-mesenchymal transition, and oral and lung SCC invasion. Whereas oral fibroblasts transfer sphingomyelins, which activate the ceramide-sphingosine-1-phosphate-STAT3 pathway and promote oral SCC invasion, lung fibroblasts transfer triglycerides to lung SCCs, thereby triggering cholesterol synthesis and invasion, which is associated with poor survival. By contrast, dermal fibroblasts are lipid poor, and cutaneous SCC is less invasive. Our data indicate that targeting fibroblast lipid synthesis and SCC lipid uptake or breakdown inhibits oral and lung epithelial cancer invasion.
    DOI:  https://doi.org/10.1038/s42255-026-01514-y
  10. Cancer Med. 2026 May;15(5): e71853
      Sidedness influences colorectal cancer (CRC) prognosis and treatment response, yet the mechanism dictating differential EGFR inhibitor (EGFRI) sensitivity is unclear. This study investigated the tumor microenvironment (TME) in relation to EGFRI eligibility-clinically defined by factors such as tumor sidedness (e.g., left-sided), RAS/BRAF wild-type status, and microsatellite stability (MSS)-using integrated single-cell RNA sequencing (scRNA-seq), with bulk RNA-seq and spatial transcriptomics validation. We found cancer cell features reflected EGFRI eligibility more strongly than sidedness. EGFRI eligible tumors exhibited high Epiregulin (EREG) expression by cancer cells. Cell interaction analysis revealed a specific "EREG/EGFR/CSF axis" in EGFRI eligible CRC: EREG derived from cancer cell stimulates EGFR-expressing non-myCAF subtypes of cancer-associated fibroblasts (CAFs), which signal via CSF to M1/M2-like Tumor-Associated Macrophages/Monocytes (TAM/TAMo), potentially promoting M2 polarization. Spatial analysis confirmed the proximity of these interacting cell populations and localized EGFR pathway activation near cancer cells specifically in eligible tumors. This study provides a TME-centric view of EGFRI eligibility, identifying a key intercellular communication network driving differential responses. These findings suggest TME features could offer more precise patient stratification than sidedness alone, potentially improving CRC therapeutic strategies.
    Keywords:  EGFR inhibitor eligibility; Epiregulin (EREG); cell–cell interaction; colorectal cancer; tumor microenvironment
    DOI:  https://doi.org/10.1002/cam4.71853
  11. Oncogene. 2026 Apr 28.
      Colorectal cancer (CRC) is one of the most commonly diagnosed and globally spread malignant diseases. Cancer-associated fibroblasts (CAFs) are key architects of the tumor microenvironment, yet their origin, stability, and interconvertibility remain poorly understood. Using transcriptomic profiling of fibroblasts from colorectal cancer (CRC) patients, we identify highly expressed (HEX) markers that define fibroblast subpopulations and uncover mechanisms governing their plasticity. We find that ADH1B marks normal colon-associated fibroblasts (NAFs), which consist of PI16-NAFs and ADAMDEC1-NAFs. ITGA3 delineates the total CAF population, which comprises myofibroblastic CAFs (myCAFs), whose characterizing markers were associated with poor prognosis and proteolytic inflammatory CAFs (piCAFs), characterized by markers not associated with prognosis. An AGT/TGM2-expressing fibroblast subset is present in both healthy and tumor tissues, suggesting alternative trajectories to the classical NAF-to-CAF transition model. While PI16-NAFs, AGT/TGM2-fibroblasts, and myCAFs maintain stable identities in long-term culture, the ADAMDEC1-NAF and piCAF phenotypes are lost in vitro. ITGA3-CAFs demonstrate dynamic plasticity, with TGF-β stably inducing myCAF formation and TNF-α or inhibition of DNA methylation promoting transient piCAF emergence. These findings redefine fibroblast heterogeneity in CRC and reveal a coexisting stable and plastic fibroblast network that may be amenable to modulation and provides a framework for future functional and translational studies. We identified highly expressed markers (HEX markers) to distinguish CAFs, NAFs and corresponding subpopulations in CRC. ADH1B characterized NAFs, which consisted of stable (solid outline) PI16-NAFs and unstable (dashed outline) ADAMDEC1-NAFs. ITGA3 identified CAFs consisting of stable myCAFs associated with poor prognosis and unstable piCAFs not associated with prognosis. AGT/TGM2 fibroblasts did not express ADH1B or ITGA3, were stable in culture and could be detected in both healthy colon and CRC. Treatment of PI16-NAFs with LPS or IFN-γ induced ADAMDEC1-NAFs, TGF-β the formation of myCAFs, while treatment with TNF-α led to the formation of piCAFs. Reduced DNA methylation converted myCAFs and PI16-NAFs into piCAFs.
    DOI:  https://doi.org/10.1038/s41388-026-03809-6
  12. bioRxiv. 2026 Apr 13. pii: 2022.02.03.479045. [Epub ahead of print]
      Drug-tolerant persisters (DTPs) represent a major obstacle to durable responses in targeted cancer therapy. DTPs are commonly described as distinct single-cell states that survive drug treatment via reversible, non-genetic mechanisms and drive tumor recurrence. Recent work demonstrates that multiple DTPs can coexist, reflecting diversity in lineage, signaling programs, or stress responses. However, each DTP is still generally viewed as a uniform cellular phenotype. Building on our prior work describing a population-level DTP termed "idling" [Paudel et al., Biophys. J. (2018) 114, 1499-1511], here we present evidence supporting a fundamentally different view: that DTPs are not single-cell states, but rather heterogeneous populations composed of multiple sub-states with distinct division and death rates that balance to produce near-zero net population growth. Using single-cell transcriptomics and lineage barcoding, we identify multiple phenotypic states within idling DTP populations, with reduced heterogeneity compared to untreated populations, and find that idling DTP cells emerge from nearly all lineages. Transcriptomic and functional analyses further reveal altered ion-channel activity in idling DTPs, which we confirm experimentally. Moreover, drug-response assays reveal increased susceptibility of idling DTPs to ferroptosis, a non-apoptotic form of regulated cell death, indicating the emergence of vulnerabilities associated with drug tolerance. Altogether, our results support a population-level view of tumor drug tolerance in which DTPs comprise stable collections of phenotypic states, shaped by treatment-defined phenotypic landscapes, which are potentially vulnerable to subsequent interventions. This perspective implies that eradicating DTPs will require a fundamental shift away from cell-type-centric strategies toward sequential treatments that progressively reduce phenotypic heterogeneity by modulating the molecular and cellular processes that establish the DTP landscape, an approach previously termed "targeted landscaping."
    DOI:  https://doi.org/10.1101/2022.02.03.479045
  13. STAR Protoc. 2026 Apr 27. pii: S2666-1667(26)00177-2. [Epub ahead of print]7(2): 104524
      Intestinal organoids (IOs) are versatile, physiologically relevant models of the human gut. In cystic fibrosis research, IOs help evaluate CFTR correctors, potentiators, and translational readthrough compounds. Here, we present a protocol for generating induced pluripotent stem cell (iPSC)-derived IOs with high phenotypic similarity to primary material. We describe steps for establishing iPSC culture, generating intestinal progenitors via directed differentiation, and maturing these intestinal progenitors using three-dimensional culture. We then detail procedures for establishing intestinal organoid cultures from these progenitors using dissection-based techniques.
    Keywords:  Cell Differentiation; Cell culture; Cell-based Assays; Organoids
    DOI:  https://doi.org/10.1016/j.xpro.2026.104524
  14. bioRxiv. 2026 Apr 15. pii: 2026.04.12.717781. [Epub ahead of print]
      Metastasis remains the primary cause of cancer-related morbidity and mortality, despite significant advances in targeted therapies. Although metastatic dissemination requires tumor cells to escape the primary lesion and colonize distant organs, the mechanisms by which primary tumor cells gain metastatic competence remain poorly understood. Increasing evidence demonstrates that fusion of tumor (i.e., neoplastic) and immune (e.g., macrophages) cells generate a distinct population of tumor-immune hybrid cells with enhanced functional ability to migrate and disseminate into peripheral blood. Herein, our study investigates tumor-macrophage hybrid cells, an underexplored population of disseminated tumor cells, and their inherent heterogeneity and acquisition of molecular mechanisms underlying their dissemination as metastatic effectors in colorectal cancer (CRC). Through hybrid cell phenotyping utilizing integrative single-cell RNA sequencing (scRNA-seq), cyclic immunofluorescence (cyCIF) and functional assays with an in vitro model of CRC hybrid cells, we identify Runt-related transcription factor 1 ( Runx1) as a central regulator of hybrid cell motility and invasion. Runx1 depletion in hybrid cells suppressed functional protease expression, chemotactic activity and extracellular matrix (ECM) invasion. Furthermore, pharmacologic inhibition of RUNX1 in an in vivo model reduced hybrid tumor growth and dissemination into peripheral blood, key attributes of metastatic spread of disease. In patients with CRC, RUNX1 + hybrid cells were identified in both primary tumor and peripheral blood, where circulating hybrid cells (CHCs) exhibited enriched migratory and epithelial-to-mesenchymal transition (EMT) phenotypes. Taken together, these findings reveal a mechanistic role for RUNX1 in driving invasive behavior of tumor-immune hybrids and highlight disseminated CHCs as an under-recognized contributor to metastatic spread and a promising noninvasive biomarker for tumor progression.
    DOI:  https://doi.org/10.64898/2026.04.12.717781
  15. Cancer Cell. 2026 Apr 30. pii: S1535-6108(26)00178-9. [Epub ahead of print]
      In this issue of Cancer Cell, Hayward et al. show that fibrotic tissue tension creates a mechanically organized mutagenic niche. A stiff stroma activates epithelial STAT3, recruits macrophages, and drives NOX-dependent lipid peroxidation, generating diffusible aldehydes that damage epithelial DNA in fibrotic tumors and mammographically dense breast tissue.
    DOI:  https://doi.org/10.1016/j.ccell.2026.04.001
  16. Clin Exp Metastasis. 2026 Apr 27. pii: 22. [Epub ahead of print]43(3):
      
    Keywords:  BMs; Colorectal cancer; FOXM1; Lipid metabolism; Metabolic adaptation
    DOI:  https://doi.org/10.1007/s10585-026-10397-y
  17. Cells Tissues Organs. 2026 Apr 30. 1-28
       BACKGROUND: Plasticity defines the capacity to change identity in response to stimuli and represents a property intrinsically inherited from embryonic development. The homeostatic epithelial turnover of many organs is guaranteed by plasticity events where stem cell progenitors generate mature differentiated epithelial cells. Plasticity is also a major hallmark of cancer cells particularly with respect to their ability to escape treatment sensitivity and adapt to harsh microenvironment.
    SUMMARY: In the context of tissue injury, modifications of epithelial cell identity represent a fundamental hallmark of damage-induced repair and regenerative processes. Epithelial reprogramming may occur through activation of dedifferentiation programs including fetal reversion and paligenosis, as well as through switches in lineage commitment, such as in transdetermination and transcommitment. Additional mechanisms rely on trans-differentiation processes between different mature epithelial cell types without the involvement of adult stem cells. The activation of all these plastic events supporting epithelial regeneration occurs not only at the phenotypic level but importantly involve epigenetic control, metabolic rearrangements and external sensing.
    KEY MESSAGES: Here we present and discuss the most recent evidences highlighting the multiple aspects of epithelial cell plasticity involved in the attempt to support repair and regenerative processes. This review will contribute to better define the broad term of plasticity, especially as emerged from latest investigations employing single-cell based omics, and identify universal and organ-specific mechanisms of injury-induced epithelial reprogramming.
    DOI:  https://doi.org/10.1159/000552260
  18. Curr Opin Genet Dev. 2026 Apr 24. pii: S0959-437X(26)00044-4. [Epub ahead of print]99 102477
      Epithelial plasticity allows committed cells to bypass rigid differentiation hierarchies, enabling efficient tissue repair through the reactivation of developmental-like programmes. In this review, we focus on the transcription factor SOX9 as a central regulator of epithelial cell fate rewiring. Essential during epithelial development and tissue morphogenesis, SOX9 is dynamically regulated across diverse epithelial tissues following injury, conferring SOX9-expressing cells with an increased 'stemness' and repair/regenerative capacity. Emerging evidence suggests that SOX9 may function as a molecular integrator of microenvironmental inputs during tissue perturbations. However, dysregulation or persistent activation of this programme carries inherent risks of fibrosis and malignancy. Future work aimed at understanding how SOX9 integrates biochemical and mechanical cues will be vital for developing strategies to harness the plastic potential of epithelial cells for regenerative medicine and prevent pathologies associated with this plasticity.
    DOI:  https://doi.org/10.1016/j.gde.2026.102477