Affinage

LAYN

Layilin · UniProt Q6UX15

Length
382 aa
Mass
43.1 kDa
Annotated
2026-06-10
39 papers in source corpus 12 papers cited in narrative 12 extracted findings
Cross-family judge faithfulness: 5/5 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

Layilin (LAYN) is a transmembrane hyaluronan (HA) receptor that couples extracellular matrix sensing to integrin activation and cytoskeletal organization, governing cell adhesion, motility, and immune effector function (PMID:32539073, PMID:22722918, PMID:39286102). In CD8+ T cells, LAYN colocalizes with integrin αLβ2 (LFA-1) and cross-linking drives LFA-1 into its activated extended state, so that LAYN loss attenuates LFA-1-dependent adhesion, reduces intratumoral T cell accumulation, and impairs direct tumor cell killing (PMID:32539073); in regulatory T cells, activation-induced LAYN limits dynamic motility to anchor Tregs within tissue (PMID:34470859). Intracellularly, LAYN interacts with talin-1 (TLN1) to drive β1 integrin signaling through c-Jun, p38 MAPK, and ERK phosphorylation, promoting EMT and cancer metastasis, with its expression controlled transcriptionally by CREB1 (PMID:39286102) and NR4A1 (PMID:36260222). As an HA receptor, LAYN transduces matrix cues bidirectionally: HA signaling through LAYN represses IL-1β-induced MMP production in chondrocytes (PMID:22722918), whereas HA oligosaccharides binding the 60–68 amino acid region of its extracellular domain activate NF-κB-dependent CCL20 secretion and pro-metastatic macrophage recruitment (PMID:36805871). LAYN additionally organizes the apical actin cap required for multiciliogenesis (PMID:37102425) and acts as a downstream effector of the p53-p21 and p16-pRB pathways in cellular senescence (PMID:21740549).

Mechanistic history

Synthesis pass · year-by-year structured walk · 12 steps
  1. 2011 Medium

    Established LAYN as a functional output of tumor-suppressor pathways rather than a passive marker, by showing it is required to execute senescence.

    Evidence RNAi screen and lentiviral shRNAmir knockdown in conditionally immortalized human fibroblasts with senescence-bypass readout

    PMID:21740549

    Open questions at the time
    • Mechanism linking LAYN to p53-p21/p16-pRB outputs not defined
    • No molecular partners identified in the senescence context
  2. 2012 Medium

    Identified LAYN as a hyaluronan receptor mediating HA's chondroprotective signaling, connecting matrix sensing to suppression of matrix-degrading enzymes.

    Evidence siRNA knockdown in human articular chondrocytes with ELISA readout of IL-1β-induced MMP-1/MMP-13

    PMID:22722918

    Open questions at the time
    • Downstream signaling between LAYN and MMP repression not mapped
    • Direct HA binding not biochemically demonstrated here
  3. 2014 Medium

    Showed LAYN extracellular engagement actively drives inflammatory mediator secretion, framing it as a signaling receptor and not merely an adhesion molecule.

    Evidence Anti-LAYN antibody cross-linking of chondrosarcoma/chondrocyte extracellular domain with ELISA for IL-8 and C5/C5a

    PMID:25150153

    Open questions at the time
    • Intracellular signaling pathway not defined
    • Cross-linking antibody may not reflect physiological ligand engagement
  4. 2015 Medium

    Linked LAYN to cytokine-specific epithelial plasticity, showing it is selectively required for TNF-α- but not TGF-β-induced EMT.

    Evidence siRNA knockdown in KMRC-1 renal tubular cells with EMT marker readouts and in vivo TNF-α administration

    PMID:26410531

    Open questions at the time
    • How TNF-α selectively engages LAYN signaling unknown
    • Downstream EMT effectors of LAYN not identified
  5. 2015 Low

    Implicated LAYN in the suppressive function of human Tregs through gain- and loss-of-function in suppression assays.

    Evidence siRNA knockdown in Tregs and overexpression in Tconvs with in vitro suppression assay

    PMID:26686412

    Open questions at the time
    • Single functional assay without mechanistic dissection
    • Effect on suppression not reconciled with later in vivo Treg findings
  6. 2020 High

    Defined the core molecular mechanism: LAYN colocalizes with and activates LFA-1 to drive T cell adhesion and antitumor cytotoxicity.

    Evidence Colocalization microscopy, cross-linking, integrin activation assay, CRISPR editing in human CD8+ T cells, and conditional knockout mice with tumor growth and ex vivo killing readouts

    PMID:32539073

    Open questions at the time
    • Direct biochemical coupling between LAYN cytoplasmic tail and integrin machinery not resolved
    • Ligand driving LFA-1 activation in vivo unspecified
  7. 2021 High

    Resolved LAYN's role in Tregs as a tissue-anchoring/motility-limiting factor distinct from suppressive capacity, revealing a counterintuitive antitumor consequence of its deletion.

    Evidence Treg-specific conditional knockout mice with live in vivo motility imaging and in vitro suppression assays

    PMID:34470859

    Open questions at the time
    • Molecular basis of motility restriction not defined
    • Tissue ligand anchoring Tregs unidentified
  8. 2021 Low

    Extended LAYN's HA-receptor adhesion role to tumor-stroma interactions in breast cancer.

    Evidence Co-culture adhesion assay between MDA-MB-231 cells and bmMSCs with HA matrix manipulation

    PMID:34707175

    Open questions at the time
    • Single adhesion assay without mechanistic dissection
    • No demonstration of downstream signaling
  9. 2022 Low

    Placed LAYN under NR4A1 transcriptional control and linked its expression to suppression of CD8+ T cell killing, with therapeutic modulation by anti-VEGFR2.

    Evidence LAYN overexpression in CD8+ T cells, co-culture killing assay, in vivo anti-VEGFR2 treatment, NR4A1 analysis

    PMID:36260222

    Open questions at the time
    • NR4A1-LAYN regulatory link not mechanistically dissected
    • Overexpression phenotype apparently opposite to knockout data, unreconciled
  10. 2023 Medium

    Mapped a discrete HA-oligosaccharide binding region on LAYN and connected its activation to NF-κB-driven chemokine secretion and pro-metastatic macrophage recruitment.

    Evidence Domain mapping/binding assays, NF-κB reporter, CCL20 ELISA, blocking antibody, and in vitro/in vivo metastasis assays in colorectal cancer

    PMID:36805871

    Open questions at the time
    • Structural basis of oHA binding not resolved
    • Receptor proximal signaling to NF-κB not detailed
  11. 2023 Medium

    Revealed a cytoskeletal/morphogenetic function: LAYN organizes the apical actin cap required for multiciliated cell formation.

    Evidence Single-cell RNA-seq, super-resolution microscopy localizing LAYN to motile cilia, and siRNA knockdown impairing multiciliogenesis across human, mouse, and pig models

    PMID:37102425

    Open questions at the time
    • Molecular link between LAYN and apical actin assembly not defined
    • Whether HA binding is involved in this role unknown
  12. 2024 Medium

    Provided the intracellular adaptor mechanism: LAYN binds talin-1 to drive β1 integrin/MAPK signaling and metastasis, under CREB1 transcriptional control.

    Evidence ChIP for CREB1 promoter binding, Co-IP for LAYN-TLN1, western blots of MAPK phosphorylation, knockdown/overexpression lines, and a nude mouse tail-vein metastasis model in cholangiocarcinoma

    PMID:39286102

    Open questions at the time
    • LAYN-TLN1 interaction shown by single Co-IP without reciprocal/structural validation
    • Direct versus indirect nature of the interaction unresolved

Open questions

Synthesis pass · forward-looking unresolved questions
  • How LAYN cytoplasmic engagement of talin/integrin machinery is regulated and reconciled across opposing pro- and anti-tumor functions in different cell types remains unresolved.
  • No structural model of the LAYN cytoplasmic tail-talin-integrin interface
  • Context-dependent switch between adhesion-promoting and motility-limiting outputs not explained
  • Physiological in vivo HA ligands for each role not defined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0008092 cytoskeletal protein binding 2 GO:0060089 molecular transducer activity 2 GO:0098631 cell adhesion mediator activity 2 GO:0060090 molecular adaptor activity 1
Localization
GO:0005856 cytoskeleton 1 GO:0005886 plasma membrane 1 GO:0005929 cilium 1
Pathway
R-HSA-162582 Signal Transduction 2 R-HSA-1643685 Disease 2 R-HSA-168256 Immune System 2 R-HSA-1474244 Extracellular matrix organization 1

Evidence

Reading pass · 12 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2020 Layilin (LAYN) colocalizes with integrin αLβ2 (LFA-1) on CD8+ T cells, and cross-linking layilin promotes the activated/extended state of LFA-1. LAYN deletion results in attenuated LFA-1-dependent cellular adhesion, and lineage-specific deletion of layilin on murine CD8+ T cells reduced their accumulation in tumors and increased tumor growth in vivo. Gene editing of LAYN in human CD8+ T cells reduced direct tumor cell killing ex vivo. Colocalization microscopy, cross-linking experiments, integrin activation assay, CRISPR gene editing in human T cells, conditional knockout in mice with tumor growth readout The Journal of experimental medicine High 32539073
2021 Layilin expression on Tregs is induced by TCR-mediated activation in the presence of IL-2 or TGF-β. In vivo, layilin expression on Tregs exerts a cumulative anchoring/motility-limiting effect, reducing Treg dynamic motility in tissue. Conditional deletion of layilin in Tregs reduced their accumulation in tumors but paradoxically enhanced immune regulation (increased tumor growth), without affecting Treg activation or suppressive capacity in vitro. Conditional knockout mice (Treg-specific LAYN deletion), live in vivo motility imaging, in vitro Treg suppression assay, flow cytometry, TCR activation experiments with cytokine stimulation Journal of immunology (Baltimore, Md. : 1950) High 34470859
2015 Layilin (LAYN) is involved in TNF-α-induced epithelial-mesenchymal transformation (EMT) of renal tubular epithelial cells. TNF-α (but not TGF-β) upregulates LAYN expression, and knockdown of LAYN in KMRC-1 cells prevents TNF-α-induced vimentin upregulation, E-cadherin downregulation, and fibroblast-like morphological change. TGF-β-induced EMT was not affected by LAYN knockdown, indicating specificity to the TNF-α pathway. siRNA knockdown, qPCR, western blotting, immunocytochemistry, in vivo TNF-α administration in mice Biochemical and biophysical research communications Medium 26410531
2014 Antibody-mediated cross-linking of the extracellular domain of layilin (LAYN) on chondrosarcoma cells enhances secretion of inflammatory factors IL-8 and complement C5/C5a. TNF-α upregulates LAYN expression in human articular chondrocytes. These findings establish LAYN as a receptor whose signaling promotes inflammatory mediator secretion in chondrocytes. Anti-LAYN antibody stimulation of extracellular domain, ELISA for cytokine secretion, qPCR, western blotting Biochemical and biophysical research communications Medium 25150153
2012 LAYN functions as a hyaluronan (HA) receptor in human articular chondrocytes. HA represses IL-1β-induced MMP-1 and MMP-13 production in chondrocytes, and this effect is significantly abrogated when LAYN is knocked down by siRNA, placing LAYN as a mediator of HA's anti-inflammatory/chondroprotective signaling. IL-1β suppresses LAYN expression in chondrocytes and synoviocytes. siRNA knockdown, ELISA for MMP-1 and MMP-13, IL-1β and TNFα stimulation experiments, in vitro cell culture Modern rheumatology Medium 22722918
2023 In colorectal cancer cells, hyaluronan oligosaccharides (oHA) activate LAYN by binding to the 60–68th amino acid region of its extracellular segment. oHA-induced LAYN activation promotes cancer cell metastasis and CCL20 secretion through the NF-κB pathway. A blocking antibody targeting LAYN prevents oHA-mediated tumor metastasis, tumor-associated macrophage (TAM) infiltration, and M2 macrophage polarization in vivo. Domain mapping (extracellular segment mutagenesis/binding assay), NF-κB pathway reporter, CCL20 ELISA, blocking antibody experiments, in vitro and in vivo metastasis assays Matrix biology : journal of the International Society for Matrix Biology Medium 36805871
2024 CREB1 binds to the LAYN promoter (validated by chromatin immunoprecipitation) and transcriptionally regulates LAYN expression. LAYN interacts with TLN1 (talin-1) as demonstrated by co-immunoprecipitation. The LAYN-TLN1 axis promotes β1 integrin signaling, including phosphorylation of c-Jun, p38 MAPK, and ERK, driving cholangiocarcinoma metastasis. Knockdown of LAYN suppressed EMT and metastasis in vivo in a nude mouse model. Chromatin immunoprecipitation (ChIP), co-immunoprecipitation (Co-IP), western blot, KD/OE cell lines, CCK-8 and Transwell assays, nude mouse tail vein metastasis model, immunofluorescence Heliyon Medium 39286102
2011 LAYN was identified as a downstream effector of the p53-p21 and p16-pRB tumour suppressor pathways in mediating cellular senescence. Direct silencing of LAYN using lentiviral shRNAmirs bypassed senescence in conditionally immortalised human fibroblasts in which these pathways were activated. RNA interference screen, lentiviral shRNAmir knockdown, microarray expression profiling, conditional senescence induction system BMC genomics Medium 21740549
2015 Knockdown of LAYN in human Tregs or overexpression in conventional T cells (Tconvs) affects Treg suppression activity in an in vitro human Treg suppression assay, implicating LAYN in the suppressive function of Tregs. siRNA knockdown in human Tregs, overexpression in Tconvs, in vitro suppression assay Human immunology Low 26686412
2022 LAYN expression on CD8+ T cells is transcriptionally regulated by the upstream transcription factor NR4A1. Overexpression of LAYN in CD8+ T cells inhibited their killing function against tumor cells in co-culture. Low-dose anti-VEGFR2 antibody treatment downregulated LAYN expression on tumor-infiltrating CD8+ T cells, correlating with enhanced anti-tumor immunity. CD8+ T cell overexpression of LAYN, co-culture tumor killing assay, flow cytometry, in vivo anti-VEGFR2 treatment, NR4A1 transcription factor analysis Cellular oncology (Dordrecht, Netherlands) Low 36260222
2023 Layilin/LAYN protein is expressed in the apical membrane regions or throughout motile cilia of multiciliated cells (MCCs), but is absent from primary cilia. LAYN silencing altered apical actin cap formation and multiciliogenesis in human, mouse, and pig MCC models, indicating a functional role in MCC formation. Single-cell RNA-seq, super-resolution microscopy, siRNA knockdown with multiciliogenesis phenotype readout FEBS letters Medium 37102425
2021 Adhesion of invasive breast cancer cells (MDA-MB-231) to bone marrow-derived mesenchymal stem cells (bmMSCs) is facilitated by a tumor-cell-induced HA-rich matrix and is mediated by the HA receptor LAYN, as demonstrated in co-culture adhesion experiments. Co-culture adhesion assay, HA matrix manipulation, identification of LAYN as mediator of adhesion Scientific reports Low 34707175

Source papers

Stage 0 corpus · 39 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2016 Transcriptional Landscape of Human Tissue Lymphocytes Unveils Uniqueness of Tumor-Infiltrating T Regulatory Cells. Immunity 542 27851914
2018 Anti-Inflammatory Effects of Intra-Articular Hyaluronic Acid: A Systematic Review. Cartilage 116 29429372
2015 Transcriptome profiling of human FoxP3+ regulatory T cells. Human immunology 79 26686412
2023 Evaluation of circulating plasma proteins in breast cancer using Mendelian randomisation. Nature communications 46 37996402
2020 Layilin augments integrin activation to promote antitumor immunity. The Journal of experimental medicine 45 32539073
2022 Integrated Analysis Reveals the Gut Microbial Metabolite TMAO Promotes Inflammatory Hepatocellular Carcinoma by Upregulating POSTN. Frontiers in cell and developmental biology 37 35676936
2011 An RNA interference screen for identifying downstream effectors of the p53 and pRB tumour suppressor pathways involved in senescence. BMC genomics 35 21740549
2014 Genome-wide unmasking of epigenetically silenced genes in lung adenocarcinoma from smokers and never smokers. Carcinogenesis 33 24398667
2015 Roles of layilin in TNF-α-induced epithelial-mesenchymal transformation of renal tubular epithelial cells. Biochemical and biophysical research communications 32 26410531
2020 Human Tregs at the materno-fetal interface show site-specific adaptation reminiscent of tumor Tregs. JCI insight 30 32809975
2024 Distinctive multicellular immunosuppressive hubs confer different intervention strategies for left- and right-sided colon cancers. Cell reports. Medicine 23 38806057
2019 Understanding tumor-infiltrating lymphocytes by single cell RNA sequencing. Advances in immunology 23 31699218
2023 Targeting LAYN inhibits colorectal cancer metastasis and tumor-associated macrophage infiltration induced by hyaluronan oligosaccharides. Matrix biology : journal of the International Society for Matrix Biology 22 36805871
2021 Layilin Anchors Regulatory T Cells in Skin. Journal of immunology (Baltimore, Md. : 1950) 20 34470859
2014 Secretion of inflammatory factors from chondrocytes by layilin signaling. Biochemical and biophysical research communications 17 25150153
2024 Mendelian randomization reveals potential causal relationships between cellular senescence-related genes and multiple cancer risks. Communications biology 16 39215079
2022 Low-dose anti-VEGFR2 therapy promotes anti-tumor immunity in lung adenocarcinoma by down-regulating the expression of layilin on tumor-infiltrating CD8+T cells. Cellular oncology (Dordrecht, Netherlands) 15 36260222
2012 Layilin, a talin-binding hyaluronan receptor, is expressed in human articular chondrocytes and synoviocytes and is down-regulated by interleukin-1β. Modern rheumatology 12 22722918
2024 A Multi-Omics Analysis of an Exhausted T Cells' Molecular Signature in Pan-Cancer. Journal of personalized medicine 9 39064019
2025 Exploring Potential Drug Targets in Multiple Cardiovascular Diseases: A Study Based on Proteome-Wide Mendelian Randomization and Colocalization Analysis. Cardiovascular therapeutics 7 40026415
2024 Prostaglandin D2 receptor 2 downstream signaling and modulation of type 2 innate lymphoid cells from patients with asthma. PloS one 7 39052598
2021 Exposure to Tobacco Smoking Induces a subset of Activated Tumor-resident Tregs in Non-Small Cell Lung Cancer. Translational oncology 7 34768099
2024 Identification of potential drug targets for allergic diseases from a genetic perspective: A mendelian randomization study. Clinical and translational allergy 6 38573314
2021 The regulatory effect of hyaluronan on human mesenchymal stem cells' fate modulates their interaction with cancer cells in vitro. Scientific reports 6 34707175
2025 The profiles of immunosuppressive microenvironment in the Lauren intestinal-type gastric adenocarcinoma. Cancer immunology, immunotherapy : CII 5 39891785
2025 Potential drug targets for asthma identified through mendelian randomization analysis. Respiratory research 3 39806440
2024 Neurological-related proteomic profiling in plasma of children with metabolic healthy and unhealthy overweight/obesity. Pediatric obesity 3 39075931
2023 The MIR34B/C genomic region contains multiple potential regulators of multiciliogenesis. FEBS letters 3 37102425
2025 Characterization of Exhausted T Cell Signatures in Pan-Cancer Settings. International journal of molecular sciences 2 40076932
2024 Associations between multiple neurological biomarkers and distal sensorimotor polyneuropathy: KORA F4/FF4 study. Diabetes/metabolism research and reviews 2 38872492
2024 Eleven neurology-related proteins measured in serum are positively correlated to the severity of diabetic neuropathy. Scientific reports 2 39048581
2024 CREB1 promotes cholangiocarcinoma metastasis through transcriptional regulation of the LAYN-mediated TLN1/β1 integrin axis: CREB1 promotes cholangiocarcinoma metastasis through regulating LAYN/TLN1/β1 integrin axis. Heliyon 2 39286102
2024 Evaluation of circulating plasma proteins in prostate cancer using mendelian randomization. Discover oncology 2 39287922
2023 Elevated Layilin-Positive Monocyte Levels in the Peripheral Blood of Patients with Systemic Lupus Erythematosus Reflect Their Autoimmune Status. Immunological investigations 2 37642473
2025 Association of the pri-miR-34b/c rs4938723 T > C polymorphism with hepatoblastoma susceptibility in Eastern Chinese children: A five-center case-control study. BMC medical genomics 1 40598293
2026 Dulaglutide Effect on Proteins Associated With CKD Progression. Kidney international reports 0 41732754
2026 Emerging biomarkers in IgA nephropathy, membranous nephropathy, and lupus nephritis. Clinica chimica acta; international journal of clinical chemistry 0 42107815
2025 Effects of Urtica dioica agglutinin on the expression of hyaluronan synthase and cell surface hyaluronic acid receptor genes. 3 Biotech 0 41376891
2025 Layilin: a multifunctional hyaluronan receptor in physiology and pathology. Frontiers in oncology 0 41404063

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