Affinage

LOXL1

Lysyl oxidase homolog 1 · UniProt Q08397

Length
574 aa
Mass
63.1 kDa
Annotated
2026-06-10
100 papers in source corpus 27 papers cited in narrative 27 extracted findings
Cross-family judge vs UniProt: tie faithfulness: 6/6 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

LOXL1 is a lysyl oxidase family enzyme that crosslinks elastin and collagen in the extracellular matrix, governing the formation and stability of elastic fiber networks across connective tissues (PMID:18265927, PMID:29366776). Loss of LOXL1 produces global connective-tissue defects: knockout mice show mechanically weaker vaginal and pelvic support tissues, pelvic organ prolapse with disorganized collagen and elastic fibers, and compromised ocular anterior-segment integrity with decreased elastin in iris and ciliary body and breakdown of the blood-aqueous barrier (PMID:18265927, PMID:24425853, PMID:34923484). In the eye, LOXL1 is a direct structural component of pathological pseudoexfoliation/exfoliation deposits, co-localizing with elastin, fibrillin-1, and fibulin-4 (PMID:19442659, PMID:25275906), and its expression is tightly controlled by disease-associated regulatory variants and pathways, including cis-acting modulation of RXRα binding and alternative splicing, a downstream retinoic acid/STRA6 axis, and promoter CpG hypermethylation (PMID:26348632, PMID:28534485, PMID:30986821). In the liver, LOXL1 crosslinks elastin specifically at the cirrhosis stage, and hepatic stellate cell-specific deletion prevents fibrosis; mechanistically LOXL1 sits upstream of TGF-β1/Smad2/3 signaling and drives a FAK/PI3K/AKT/HIF1α cascade in which HIF1α directly binds the LOXL1 promoter to create a positive feedback loop (PMID:29366776, PMID:30257391, PMID:34569648, PMID:34014450). Beyond its canonical ECM role, LOXL1 modulates intracellular signaling in cancer with context-dependent outcomes: it antagonizes Ras/ERK and suppresses YAP through MST1/2-dependent Hippo activation in a tumor-suppressive capacity (PMID:17456585, PMID:32912229), while in other contexts it stabilizes the anti-apoptotic factor BAG2 by preventing its ubiquitination and promotes angiogenesis through a fibulin-5/αvβ3 integrin/FAK-MAPK axis (PMID:32424143, PMID:33614230). LOXL1 expression is further regulated by integrin signaling (ITGA8, integrin α11) and by direct targeting of its 3'-UTR by miR-15b/16 (PMID:31121900, PMID:34569648, PMID:33545201).

Mechanistic history

Synthesis pass · year-by-year structured walk · 13 steps
  1. 1999 Low

    Established the first hint that LOXL1, unlike LOX, has distinct collagen-substrate associations, raising the question of functional specialization within the lysyl oxidase family.

    Evidence Real-time RT-PCR correlating Loxl1 and procollagen mRNAs during CCl4 liver fibrosis in mice

    PMID:10022501

    Open questions at the time
    • mRNA correlation only; no direct enzymatic or protein-level confirmation of substrate specificity
    • does not demonstrate crosslinking activity on any substrate
  2. 2004 Medium

    Mapped where LOXL1 protein resides relative to LOX, showing overlapping and distinct tissue zones and unexpected nuclear localization, broadening the scope of possible LOXL1 functions beyond the ECM.

    Evidence Comparative immunohistochemistry across developing mouse tissues

    PMID:15609098

    Open questions at the time
    • nuclear function not defined
    • localization does not establish a molecular activity
  3. 2008 Medium

    Demonstrated that LOXL1 is required for the mechanical integrity of connective tissues, directly linking the enzyme to ECM crosslinking-dependent tissue strength.

    Evidence Biomechanical tensile testing of vaginal tissue in Loxl1-/- vs wild-type mice; IHC and RT-PCR across rat aorta aging

    PMID:18265927 PMID:18803461

    Open questions at the time
    • molecular crosslink substrates not biochemically quantified in these assays
    • tissue-level phenotype does not isolate elastin vs collagen contribution
  4. 2009 Medium

    Identified LOXL1 as a direct physical constituent of pseudoexfoliation deposits, connecting the crosslinking enzyme to a defined human ocular pathology.

    Evidence Mass spectrometry of surgically isolated PEX material with IHC confirmation

    PMID:19442659

    Open questions at the time
    • does not establish whether LOXL1 enzymatic activity drives deposit formation
    • causality between LOXL1 and PEX assembly not tested
  5. 2011 Medium

    Showed environmental and cytokine stimuli (TGF-β1, oxidative stress, UV, hypoxia) induce LOXL1 and PEX-like fibril assembly, and that risk haplotypes alter LOXL1 expression, linking genetic risk to a stress-responsive matrix phenotype.

    Evidence Genotype-stratified human Tenon fibroblast cultures with qPCR, Western, IHC, EM

    PMID:21948647

    Open questions at the time
    • modest expression differences between haplotypes
    • causal link from expression change to fibril phenotype not isolated
  6. 2014 High

    Defined the in vivo ocular requirement for LOXL1 in blood-aqueous barrier and elastin maintenance, and confirmed LOXL1 as a major XFS deposit component co-localizing with elastic fiber proteins.

    Evidence Loxl1 null mouse clinical/histological analysis and fluorescein angiography; human XFS tissue PCR/IHC/Western/EM

    PMID:24425853 PMID:25275906

    Open questions at the time
    • absence of IOP elevation leaves glaucoma mechanism unexplained
    • stage-dependent up- then down-regulation of LOXL1 in disease not mechanistically resolved
  7. 2015 Medium

    Localized PEX risk to non-coding regulatory elements at the LOXL1 locus, showing risk alleles modulate promoter activity and an antisense lncRNA, and that the locus is epigenetically silenced in disease tissue.

    Evidence Deep sequencing, luciferase promoter assays, multi-population replication; pyrosequencing of CpG islands in lens capsule

    PMID:26307087 PMID:26348632

    Open questions at the time
    • lncRNA mechanism on LOXL1 protein function not established
    • small sample size for methylation analysis
  8. 2016 Medium

    Demonstrated that PEX coding variants alter LOXL1 surface electrostatics and protease processing of secreted protein, providing a protein-level mechanism for genetic risk.

    Evidence Molecular modelling and immunoprecipitation of secreted LOXL1 variants from fibroblasts

    PMID:26997634

    Open questions at the time
    • BMP-1 as processing protease only inferred
    • functional consequence for crosslinking activity not measured
  9. 2017 High

    Pinpointed a causal cis-regulatory variant acting through RXRα binding and alternative splicing to reduce LOXL1 expression, establishing the molecular basis of the LOXL1 PEX association.

    Evidence GWAS with three-population replication, reporter and TF-binding assays, splicing analysis in patient tissues; cartilage stiffness correlation study

    PMID:28534485 PMID:28623328

    Open questions at the time
    • tissue-specific splicing outcomes not fully enumerated
    • cartilage stiffness link is correlative without LOXL1 manipulation
  10. 2018 High

    Established LOXL1 as a stage-specific elastin crosslinker in liver cirrhosis whose knockdown arrests disease, and placed LOXL1 upstream of TGF-β1/Smad2/3 fibrogenic signaling.

    Evidence CCl4 cirrhosis model with AAV shRNA knockdown and biochemical crosslink quantification; siRNA/overexpression epistasis in LX-2 cells

    PMID:29366776 PMID:30257391

    Open questions at the time
    • how an extracellular enzyme modulates intracellular Smad phosphorylation not mechanistically resolved
    • direct enzymatic crosslinking of elastin vs indirect signaling effects not fully separated
  11. 2019 Medium

    Showed LOXL1 functions in tumor stroma, regulated by integrins, to remodel and align collagen and support tumor growth, and that it can drive EMT in epithelial tumor cells.

    Evidence LOXL1 knockdown/overexpression in CAFs with SHG microscopy and NSCLC xenografts; LOXL1 overexpression and EMT markers in gastric cancer cells

    PMID:31121900 PMID:33095806

    Open questions at the time
    • EMT effect lacks defined mechanistic pathway
    • integrin α11 to LOXL1 regulatory link not fully dissected
  12. 2020 Medium

    Revealed two intracellular signaling roles for LOXL1 in cancer: enzymatic stabilization of BAG2 via blocking its ubiquitination (anti-apoptotic) and MST1/2-dependent Hippo activation suppressing YAP (tumor-suppressive).

    Evidence Co-IP, interaction-site modeling, ubiquitination assays, and xenografts in glioma; Co-IP with MST1/2 and phosphorylation/xenograft assays in colorectal cancer

    PMID:32424143 PMID:32912229

    Open questions at the time
    • opposing tumor-suppressive vs pro-survival roles not reconciled across tissues
    • single-lab Co-IP findings for each interaction
  13. 2021 High

    Defined a hepatic stellate cell autonomous LOXL1 fibrosis circuit (ITGA8 → FAK/PI3K/AKT/HIF1α with HIF1α-driven positive feedback), additional regulatory inputs (miR-15b/16, RA/STRA6, rs7173049), and broader roles in angiogenesis, ECM-supported ER signaling, and NASH.

    Evidence HSC-specific conditional Loxl1 knockout with RNA-seq, ChIP, and pharmacological epistasis; miRNA 3'-UTR reporter; multi-population enhancer/TF-binding assays; FBLN5 Co-IP; ILC and NASH models

    PMID:30986821 PMID:33545201 PMID:33614230 PMID:33616307 PMID:34014450 PMID:34569648 PMID:34923484

    Open questions at the time
    • integration of the many parallel signaling and regulatory axes into one model is incomplete
    • relative contribution of enzymatic crosslinking vs signaling adaptor roles unresolved

Open questions

Synthesis pass · forward-looking unresolved questions
  • How a primarily extracellular crosslinking enzyme exerts the diverse intracellular signaling effects attributed to it (Smad2/3, Hippo, BAG2 stabilization, FAK/HIF1α) and the function of its reported nuclear pool remain unresolved.
  • no structural or biochemical mechanism linking secreted LOXL1 to intracellular kinase pathways
  • nuclear LOXL1 function uncharacterized
  • no reconstituted enzymatic assay defining elastin vs collagen substrate preference

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0005198 structural molecule activity 3 GO:0016491 oxidoreductase activity 2 GO:0140096 catalytic activity, acting on a protein 1
Localization
GO:0005576 extracellular region 3 GO:0031012 extracellular matrix 3 GO:0005634 nucleus 1
Pathway
R-HSA-162582 Signal Transduction 4 R-HSA-1643685 Disease 4 R-HSA-1474244 Extracellular matrix organization 3
Partners
BAG2FBLN5MST1MST2

Evidence

Reading pass · 27 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1999 LOXL1 (mouse LOXL) mRNA is co-induced with type III procollagen mRNA during early carbon tetrachloride-induced liver fibrosis, whereas LOX co-induces with type I procollagen, suggesting LOXL1 has distinct collagen-type substrate specificity compared to LOX. Real-time RT-PCR of mRNA levels during CCl4-induced liver fibrosis in mice Journal of cellular biochemistry Low 10022501
2004 LOXL1 protein co-localizes with LOX in multiple tissues (skin, aorta, heart, lung, liver, cartilage) but occupies distinct zones in kidney, gut, retina, and gonads; both proteins are also found in cell nuclei across several tissues, suggesting nuclear as well as extracellular functions. Comparative immunohistochemistry in normal and developing mouse tissues across developmental stages Journal of molecular histology Medium 15609098
2007 LOXL1 overexpression in human bladder cancer cells decreases colony formation and antagonizes Ras-induced activation of the ERK signaling pathway, indicating a tumor-suppressive function mediated through inhibition of Ras/ERK. Re-expression of LOXL1 in bladder cancer cell lines; colony formation assay; Western blot for ERK phosphorylation Cancer research Medium 17456585
2008 Homozygous LOXL1 knockout (Loxl1−/−) mice develop mechanically weaker vaginal and pelvic supportive tissues, with a 31% decrease in ultimate load at failure, indicating a global connective tissue defect attributable to loss of LOXL1-mediated extracellular matrix crosslinking. Biomechanical tensile testing to failure of vaginal tissue complexes from Loxl1−/− and wild-type mice International urogynecology journal and pelvic floor dysfunction Medium 18265927
2008 LOXL1 expression in rat aorta is maintained throughout life predominantly in intima and media (in contrast to LOX which peaks embryonically then drops), suggesting LOXL1 plays a specific role in adult elastin and collagen fiber remodeling distinct from the developmental role of LOX. Immunohistochemistry and real-time RT-PCR across developmental, growth, and aging stages in two rat strains; combined with elastin/collagen content measurements Rejuvenation research Low 18803461
2009 LOXL1 protein is directly identified as a component of surgically isolated pseudoexfoliation (PEX) material from anterior lens capsules, consistent with its role as a cross-linking enzyme participating in formation of pathological PEX deposits. Direct mass spectrometry of surgically isolated PEX material; immunohistochemical confirmation on lens capsules Experimental eye research Medium 19442659
2011 TGF-β1, oxidative stress, UV light, and hypoxia upregulate LOXL1 expression and promote assembly of elastin/microfibrillar networks into PEX-like fibrils in human Tenon fibroblasts; cells carrying the high-risk LOXL1 haplotype show slightly decreased basal and stimulated LOXL1 expression compared to low-risk haplotype cells. Exposure of cultured human Tenon's capsule fibroblasts (high- vs. low-risk LOXL1 haplotypes) to pathogenic stimuli; quantitative real-time PCR, Western blot, immunohistochemistry, electron microscopy Investigative ophthalmology & visual science Medium 21948647
2014 Loxl1−/− mice show compromise of the blood-aqueous humor barrier (increased fluorescein diffusion into anterior chamber), anterior lens subcapsular vesiculation, and decreased elastin in iris and ciliary body, but no macromolecular deposit accumulation or elevated intraocular pressure, demonstrating that LOXL1 is required for ocular anterior segment integrity and elastin maintenance. Clinical and histological analysis, fluorescein angiography, IOP measurement, and immunohistochemistry in Loxl1 null vs. C57BL control mice Investigative ophthalmology & visual science High 24425853
2014 LOXL1 is a major component of exfoliation (XFS) material, co-localizing with elastin, fibrillin-1, and fibulin-4; LOXL1 expression is upregulated in early XFS anterior eye tissues but downregulated in advanced stages and specifically downregulated in the lamina cribrosa of XFS eyes, where ultrastructural elastic fiber network alterations are observed. Real-time PCR, immunohistochemistry, Western blot, and electron microscopy on human XFS and control eye tissues; in vitro cell stimulation assays Journal of glaucoma Medium 25275906
2015 A 7-kb region spanning introns 1-2 of LOXL1 contains an enhancer-like promoter for the antisense lncRNA LOXL1-AS1; XFS risk alleles in this region functionally modulate promoter activity, and LOXL1-AS1 expression is altered by oxidative stress and cyclic mechanical stress in ocular cells. Deep sequencing of LOXL1 locus; luciferase promoter activity assays; multi-population replication; expression analysis in stressed ocular cells Human molecular genetics Medium 26307087
2015 LOXL1 promoter CpG islands are hypermethylated in lens capsule samples from PEX patients compared to age-related cataract controls, with corresponding reduction in LOXL1 mRNA and protein, indicating epigenetic silencing of LOXL1 in PEX tissue. Pyrosequencing of CpG islands; quantitative real-time PCR and Western blot on lens capsule specimens Investigative ophthalmology & visual science Medium 26348632
2016 The PEX-associated coding variants Arg141Leu and Gly153Asp in LOXL1 affect surface electrostatics at likely protein-protein interaction sites and alter processing of secreted LOXL1 protein variants by endogenous proteases (possibly BMP-1), as shown by differential immunoprecipitation of secreted forms from fibroblasts. Molecular modelling; ectopic expression in RFL-6 rat fetal lung fibroblasts; immunoprecipitation of secreted protein variants; immunofluorescence Experimental eye research Medium 26997634
2017 The rs11638944:C>G transversion in intron 1-2 of LOXL1 exerts a cis-acting effect by (a) differentially modulating binding of transcription factor RXRα and (b) altering alternative splicing of LOXL1, leading to reduced LOXL1 mRNA levels in risk allele carriers. Genome-wide association scan; independent population replication; reporter and transcription factor binding assays; RNA splicing analysis in patient cells and tissues Nature communications High 28534485
2018 LOXL1 specifically crosslinks elastin in the liver: LOXL1 expression coincides with appearance of insoluble/crosslinked elastin only in cirrhosis (not earlier fibrosis stages); AAV-delivered shRNA knockdown of LOXL1 in cirrhotic mice reduces insoluble elastin area by 58% and collagen area by 51%, arresting disease progression. CCl4-induced liver fibrosis/cirrhosis mouse model; AAV2/8 shRNA knockdown; biochemical quantification of crosslinked elastin and collagen; immunolocalization of LOXL1 with α-SMA; in vitro LOXL1 silencing in LX-2 cells Biochimica et biophysica acta. Molecular basis of disease High 29366776
2018 LOXL1 knockdown in TGF-β1-stimulated hepatic stellate cells (LX-2) suppresses cell proliferation and fibrogenic gene expression (TIMP1, α-SMA, collagen type I) and reduces phosphorylation of Smad2 and Smad3; conversely, Smad2/3 knockdown abolishes proliferation and fibrogenesis driven by LOXL1 overexpression, placing LOXL1 upstream of TGF-β1/Smad2/3 signaling. siRNA knockdown and overexpression in LX-2 cells; Western blot for Smad2/3 phosphorylation; proliferation assays; epistasis by double knockdown Biomedicine & pharmacotherapy Medium 30257391
2019 LOXL1 expression in cancer-associated fibroblasts is regulated by integrin α11; LOXL1 promotes collagen matrix remodeling and collagen fiber alignment in vitro and in vivo, supporting NSCLC tumor growth in a xenograft model. LOXL1 knockdown and overexpression in CAFs; collagen gel contraction assays; second-harmonic generation microscopy for fiber alignment; NSCLC xenograft tumor growth Cancers Medium 31121900
2019 A non-coding variant rs7173049-G downstream of LOXL1 correlates with allele-specific binding of thyroid hormone receptor beta, increased expression of ISLR2 and STRA6, and a consistent protective effect on PEX risk across nine populations; siRNA-mediated downregulation of retinoic acid signaling (STRA6 pathway) induces upregulation of LOXL1 and PEX-associated matrix genes in ocular cells, placing LOXL1 expression downstream of the RA/STRA6 axis. Deep sequencing; luciferase enhancer assays; transcription factor binding assays; multi-population replication; siRNA knockdown of RA pathway components with LOXL1 expression readout Human molecular genetics High 30986821
2020 LOXL1 interacts with BAG2 via a hydrogen bond between LOXL1-D515 and BAG2-K186; LOXL1's lysyl oxidase enzymatic activity prevents ubiquitination of BAG2-K186, thereby stabilizing BAG2 protein and conferring antiapoptotic activity in glioma cells. LOXL1 expression is specifically upregulated through the VEGFR-Src-CEBPA transcriptional axis. Co-immunoprecipitation; structural modeling of interaction; gain/loss-of-function in glioma cells and xenografts; ubiquitination assays; pathway epistasis (VEGFR-Src-CEBPA) Cell death and differentiation Medium 32424143
2020 LOXL1 inhibits colorectal cancer progression by interacting with MST1/2 kinases and increasing their phosphorylation, thereby suppressing YAP transcriptional activity via the Hippo pathway; overexpression of LOXL1 inhibits metastasis and tumor growth in vivo. Co-immunoprecipitation of LOXL1 with MST1/2; Western blot for MST1/2 phosphorylation; colony formation, migration/invasion, and xenograft assays Cell communication and signaling Medium 32912229
2021 LOXL1 silencing in ILC (invasive lobular carcinoma) cell lines decreases tumor growth, invasion, and metastasis in intraductal xenograft models by disrupting ECM structure; this is associated with decreased estrogen receptor (ER) signaling, indicating LOXL1-mediated ECM remodeling supports ER activity in ILC. LOXL1 shRNA knockdown and pan-LOX inhibitor BAPN in ILC intraductal xenograft models; ECM structural analysis; ER signaling readouts EMBO molecular medicine Medium 33616307
2021 Hepatic stellate cell-specific LOXL1 knockout prevents CCl4-induced hepatic fibrosis and inflammation; RNA-seq identifies integrin α8 (ITGA8) as a key upstream regulator; LOXL1 activates the FAK/PI3K/AKT/HIF1α pathway, and HIF1α directly binds the LOXL1 promoter creating a positive feedback loop; FAK or PI3K inhibitors reduce LOXL1 expression. Conditional Loxl1 knockout in HSCs (Loxl1-Gfap-cre mice); RNA sequencing; siRNA silencing of ITGA8; ChIP/promoter binding assays for HIF1α; pharmacological inhibition of FAK/PI3K; fibrosis quantification (Sirius red, hydroxyproline) FASEB journal High 34569648
2021 Loxl1-deficient mice develop pelvic organ prolapse with disordered ECM (disarranged collagen and elastic fibers), urinary dysfunction, decreased mechanical properties of pelvic floor, upregulated ECM catabolism gene expression, and elevated inflammatory response, consistent with pathology in human POP patients. Histology, GO/transcriptomic analysis of differentially expressed genes, urodynamic functional testing, and mechanical testing in Loxl1 knockout mice; comparison to human prolapse tissue Aging Medium 34923484
2021 LOXL1 interacts with fibulin-5 (FBLN5), which then binds αvβ3 integrin via an RGD domain, activating FAK-MAPK signaling in vascular endothelial cells to promote angiogenesis in intrahepatic cholangiocarcinoma; LOXL1 overexpression promotes proliferation, colony formation, metastasis, and angiogenesis in vivo and in vitro. Co-immunoprecipitation of LOXL1 with FBLN5; RGD domain-dependent binding assay; Western blot for FAK-MAPK activation; xenograft and angiogenesis assays Molecular therapy. Nucleic acids Medium 33614230
2021 HSC-specific LOXL1 deficiency in a CDAA diet-induced non-obese NASH model reduces inflammation, fibrosis, and hepatic steatosis and restores adipose tissue function; serum LOXL1 positively correlates with histological fibrosis and inversely with leptin in non-obese NAFLD patients. Conditional HSC-specific Loxl1 knockout mice (Loxl1fl/fl Gfap-cre); CDAA diet NASH model; liver histology and profibrogenic/pro-inflammatory gene expression; correlation analysis in human NAFLD patients Hepatology international Medium 34014450
2017 LOXL1 gene expression is upregulated ~10-fold in platelet-rich plasma (PRP)-treated immature articular cartilage, correlating with a 5-fold increase in cartilage stiffness measured by atomic force microscopy, suggesting LOXL1-mediated ECM crosslinking contributes to PRP-induced cartilage maturation. In vitro PRP treatment of immature cartilage explants; gene expression profiling; atomic force microscopy for stiffness; BrdU incorporation for proliferation Scientific reports Low 28623328
2021 miR-15b and miR-16 directly bind the 3'-UTR of LOXL1 mRNA and suppress LOXL1 expression in hepatic stellate cells; LOXL1 knockdown attenuates the pro-fibrogenic and pro-proliferative effects of miR-15b/16 downregulation, placing LOXL1 as a downstream effector of miR-15b/16 in TGF-β1-driven fibrosis. 3'-UTR luciferase reporter assay; miRNA overexpression and knockdown in LX-2 cells; Western blot for Smad2/3 phosphorylation and fibrogenic markers; epistasis by LOXL1 knockdown in miR-15b/16-depleted cells Life sciences Medium 33545201
2019 LOXL1 overexpression in gastric cancer cells promotes epithelial-mesenchymal transition (EMT), reducing CDH1 expression while increasing VIM, CDH2, SNAI2, and PLS3, and enhancing cell migration capacity. LOXL1 overexpression in GC cell lines; Western blot and morphological analysis for EMT markers; migration assay PloS one Low 33095806

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1991 Cloning and sequencing of Lol pI, the major allergenic protein of rye-grass pollen. FEBS letters 118 2001733
2008 Molecular pathology of pseudoexfoliation syndrome/glaucoma--new insights from LOXL1 gene associations. Experimental eye research 104 18809397
2015 Outer membrane lipoprotein biogenesis: Lol is not the end. Philosophical transactions of the Royal Society of London. Series B, Biological sciences 100 26370942
2008 Association of LOXL1 common sequence variants in German and Italian patients with pseudoexfoliation syndrome and pseudoexfoliation glaucoma. Investigative ophthalmology & visual science 96 18385063
2007 LOXL1 and LOXL4 are epigenetically silenced and can inhibit ras/extracellular signal-regulated kinase signaling pathway in human bladder cancer. Cancer research 94 17456585
2018 Long noncoding RNA LOXL1-AS1 regulates prostate cancer cell proliferation and cell cycle progression through miR-541-3p and CCND1. Biochemical and biophysical research communications 88 30278884
2010 Major LOXL1 risk allele is reversed in exfoliation glaucoma in a black South African population. Molecular vision 86 20431720
2008 Association of LOXL1 gene polymorphisms with pseudoexfoliation in the Japanese. Investigative ophthalmology & visual science 84 18450598
2011 Regulation of lysyl oxidase-like 1 (LOXL1) and elastin-related genes by pathogenic factors associated with pseudoexfoliation syndrome. Investigative ophthalmology & visual science 79 21948647
2008 Evaluation of LOXL1 gene polymorphisms in exfoliation syndrome and exfoliation glaucoma. Molecular vision 79 18385788
2004 Comparative immunocytochemical localization of lysyl oxidase (LOX) and the lysyl oxidase-like (LOXL) proteins: changes in the expression of LOXL during development and growth of mouse tissues. Journal of molecular histology 78 15609098
1988 Association of HLA-DR3 with human immune response to Lol p I and Lol p II allergens in allergic subjects. Tissue antigens 77 3261461
2009 Identification of LOXL1 protein and Apolipoprotein E as components of surgically isolated pseudoexfoliation material by direct mass spectrometry. Experimental eye research 71 19442659
2018 LncRNA LOXL1-AS1 Promotes the Proliferation and Metastasis of Medulloblastoma by Activating the PI3K/AKT Pathway. Analytical cellular pathology (Amsterdam) 70 30050750
2018 Inhibition of lysyl oxidase-like 1 (LOXL1) expression arrests liver fibrosis progression in cirrhosis by reducing elastin crosslinking. Biochimica et biophysica acta. Molecular basis of disease 69 29366776
2019 LncRNA LOXL1-AS1 facilitates the tumorigenesis and stemness of gastric carcinoma via regulation of miR-708-5p/USF1 pathway. Cell proliferation 68 31468594
2009 Evaluation of LOXL1 polymorphisms in exfoliation syndrome in a Chinese population. Molecular vision 68 19936304
2008 LOXL1 genetic polymorphisms are associated with exfoliation glaucoma in the Japanese population. Molecular vision 68 18552979
2008 Evaluation of LOXL1 polymorphisms in eyes with exfoliation glaucoma in Japanese. Molecular vision 68 18648524
1989 Complete primary structure of a Lolium perenne (perennial rye grass) pollen allergen, Lol p III: comparison with known Lol p I and II sequences. Biochemistry 67 2605214
2015 Genetic variants and cellular stressors associated with exfoliation syndrome modulate promoter activity of a lncRNA within the LOXL1 locus. Human molecular genetics 65 26307087
2008 Lysyl oxidase-like protein 1 (LOXL1) gene polymorphisms and exfoliation glaucoma in a Central European population. Molecular vision 61 18483563
2019 LOXL1 Is Regulated by Integrin α11 and Promotes Non-Small Cell Lung Cancer Tumorigenicity. Cancers 58 31121900
2009 Association of LOXL1 gene with Finnish exfoliation syndrome patients. Journal of human genetics 57 19343041
2008 Differential expression of lysyl oxidases LOXL1 and LOX during growth and aging suggests specific roles in elastin and collagen fiber remodeling in rat aorta. Rejuvenation research 56 18803461
2002 Mutants of the major ryegrass pollen allergen, Lol p 5, with reduced IgE-binding capacity: candidates for grass pollen-specific immunotherapy. European journal of immunology 56 11782018
2018 Silencing LncRNA LOXL1-AS1 attenuates mesenchymal characteristics of glioblastoma via NF-κB pathway. Biochemical and biophysical research communications 55 29678575
2019 LncRNA LOXL1-AS1/miR-let-7a-5p/EGFR-related pathway regulates the doxorubicin resistance of prostate cancer DU-145 cells. IUBMB life 54 31188543
2017 Pseudoexfoliation syndrome-associated genetic variants affect transcription factor binding and alternative splicing of LOXL1. Nature communications 50 28534485
2021 Intraductal xenografts show lobular carcinoma cells rely on their own extracellular matrix and LOXL1. EMBO molecular medicine 49 33616307
2009 Association of LOXL1 polymorphisms with pseudoexfoliation in the Chinese. Molecular vision 49 19503743
2011 Outer membrane targeting of Pseudomonas aeruginosa proteins shows variable dependence on the components of Bam and Lol machineries. mBio 44 22147293
2019 LncRNA LOXL1-AS1/miR-28-5p/SEMA7A axis facilitates pancreatic cancer progression. Cell biochemistry and function 42 31732974
2020 LOXL1 confers antiapoptosis and promotes gliomagenesis through stabilizing BAG2. Cell death and differentiation 41 32424143
2014 Disruption of the blood-aqueous barrier and lens abnormalities in mice lacking lysyl oxidase-like 1 (LOXL1). Investigative ophthalmology & visual science 41 24425853
2008 Genetic association of LOXL1 gene variants and exfoliation glaucoma in a Utah cohort. Cell cycle (Georgetown, Tex.) 41 18287813
2019 Long non-coding RNA LOXL1-AS1 acts as a ceRNA for miR-324-3p to contribute to cholangiocarcinoma progression via modulation of ATP-binding cassette transporter A1. Biochemical and biophysical research communications 40 31003776
2019 The role of lysyl oxidase-like 1 (LOXL1) in exfoliation syndrome and glaucoma. Experimental eye research 40 31563608
2021 LOXL1 exerts oncogenesis and stimulates angiogenesis through the LOXL1-FBLN5/αvβ3 integrin/FAK-MAPK axis in ICC. Molecular therapy. Nucleic acids 39 33614230
2020 LncRNA LOXL1-AS1 is transcriptionally activated by JUND and contributes to osteoarthritis progression via targeting the miR-423-5p/KDM5C axis. Life sciences 39 32679142
2018 Knockdown of LOXL1 inhibits TGF-β1-induced proliferation and fibrogenesis of hepatic stellate cells by inhibition of Smad2/3 phosphorylation. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 39 30257391
2008 LOXL1 deficiency negatively impacts the biomechanical properties of the mouse vagina and supportive tissues. International urogynecology journal and pelvic floor dysfunction 39 18265927
2019 Long Non-coding RNA LOXL1-AS1 Drives Breast Cancer Invasion and Metastasis by Antagonizing miR-708-5p Expression and Activity. Molecular therapy. Nucleic acids 38 31945728
2010 Analysis of LOXL1 polymorphisms in a Saudi Arabian population with pseudoexfoliation glaucoma. Molecular vision 38 21197115
1999 Coexpression of the lysyl oxidase-like gene (LOXL) and the gene encoding type III procollagen in induced liver fibrosis. Journal of cellular biochemistry 38 10022501
1994 IgE and IgG cross-reactivity among Lol p I and Lol p II/III. Identification of the C-termini of Lol p I, II, and III as cross-reactive structures. Allergy 38 7518655
2011 An investigation into LOXL1 variants in black South African individuals with exfoliation syndrome. Archives of ophthalmology (Chicago, Ill. : 1960) 37 21320968
2020 Non-coding RNA LOXL1-AS1 exhibits oncogenic activity in ovarian cancer via regulation of miR-18b-5p/VMA21 axis. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 36 32058209
2004 Downregulation of tissue factor by RNA interference in human melanoma LOX-L cells reduces pulmonary metastasis in nude mice. International journal of cancer 36 15386437
2014 Expression and regulation of LOXL1 and elastin-related genes in eyes with exfoliation syndrome. Journal of glaucoma 34 25275906
2021 Hepatic stellate cells-specific LOXL1 deficiency abrogates hepatic inflammation, fibrosis, and corrects lipid metabolic abnormalities in non-obese NASH mice. Hepatology international 33 34014450
2020 LncRNA LOXL1-AS1 controls osteogenic and adipocytic differentiation of bone marrow mesenchymal stem cells in postmenopausal osteoporosis through regulating the miR-196a-5p/Hmga2 axis. Journal of bone and mineral metabolism 32 32651705
2019 LncRNA LOXL1-AS1 regulates the tumorigenesis and development of lung adenocarcinoma through sponging miR-423-5p and targeting MYBL2. Cancer medicine 31 31758653
2020 LncRNA LOXL1-AS1 promotes endometrial cancer progression by sponging miR-28-5p to upregulate RAP1B expression. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 30 32006897
2020 LOXL1 modulates the malignant progression of colorectal cancer by inhibiting the transcriptional activity of YAP. Cell communication and signaling : CCS 30 32912229
2015 LOXL1 Hypermethylation in Pseudoexfoliation Syndrome in the Uighur Population. Investigative ophthalmology & visual science 30 26348632
2008 The LOXL1 gene variations are not associated with primary open-angle and primary angle-closure glaucomas. Investigative ophthalmology & visual science 30 18223248
2008 Exfoliation syndrome and exfoliation glaucoma-associated LOXL1 variations are not involved in pigment dispersion syndrome and pigmentary glaucoma. Molecular vision 30 18618003
2011 Prevalence of high-risk alleles in the LOXL1 gene and its association with pseudoexfoliation syndrome and exfoliation glaucoma in a Latin American population. Ophthalmic genetics 29 21970694
2018 Exfoliation Syndrome: A Disease of Autophagy and LOXL1 Proteopathy. Journal of glaucoma 27 29547474
2016 LOXL1 Gene Polymorphism With Exfoliation Syndrome/Exfoliation Glaucoma: A Meta-Analysis. Journal of glaucoma 27 25304275
2013 Association of LOXL1 gene polymorphisms with exfoliation syndrome/glaucoma and primary open angle glaucoma in a Turkish population. Molecular vision 27 23378724
2010 LOXL1 expression in lens capsule tissue specimens from individuals with pseudoexfoliation syndrome and glaucoma. Molecular vision 27 21139690
1995 Mapping of the antigenic and allergenic epitopes of Lol p VB using gene fragmentation. Molecular immunology 27 7723775
2019 LncRNA LOXL1-AS1 inhibited cell proliferation, migration and invasion as well as induced apoptosis in breast cancer via regulating miR-143-3p. European review for medical and pharmacological sciences 26 31841194
2021 Selective depletion of hepatic stellate cells-specific LOXL1 alleviates liver fibrosis. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 25 34569648
2019 LncRNA LOXL1-AS1 promotes invasion and proliferation of non-small-cell lung cancer through targeting miR-324-3p. American journal of translational research 25 31737192
2007 Systematic screening of lysyl oxidase-like (LOXL) family genes demonstrates that LOXL2 is a susceptibility gene to intracranial aneurysms. Human genetics 25 17287949
1997 Biochemical and immunological characterization of recombinant allergen Lol p 1. European journal of biochemistry 25 9395340
2020 LOXL1-AS1/miR-515-5p/STAT3 Positive Feedback Loop Facilitates Cell Proliferation and Migration in Atherosclerosis. Journal of cardiovascular pharmacology 24 32453072
2020 LOXL1-AS1 contributes to the proliferation and migration of laryngocarcinoma cells through miR-589-5p/TRAF6 axis. Cancer cell international 24 33061856
2008 Evaluation of LOXL1 polymorphisms in primary open-angle glaucoma in southern and northern Chinese. Molecular vision 24 19098994
2021 LOXL1-AS1 communicating with TIAR modulates vasculogenic mimicry in glioma via regulation of the miR-374b-5p/MMP14 axis. Journal of cellular and molecular medicine 23 34890108
2021 Structural, functional and molecular pathogenesis of pelvic organ prolapse in patient and Loxl1 deficient mice. Aging 23 34923484
2020 Long non-conding RNA LOXL1-AS1 sponges miR-589-5p to up-regulate CBX5 expression in renal cell carcinoma. Bioscience reports 23 33185692
2016 Biological effect of LOXL1 coding variants associated with pseudoexfoliation syndrome. Experimental eye research 23 26997634
2021 MiR-15b and miR-16 suppress TGF-β1-induced proliferation and fibrogenesis by regulating LOXL1 in hepatic stellate cells. Life sciences 22 33545201
2019 Transcriptome analysis of luminal breast cancer reveals a role for LOL in tumor progression and tamoxifen resistance. International journal of cancer 22 30720865
1993 Cloning, expression, and immunological characterization of recombinant Lolium perenne allergen Lol p II. The Journal of biological chemistry 22 7691817
2021 LOXL1 gene polymorphism candidates for exfoliation glaucoma are also associated with a risk for primary open-angle glaucoma in a Caucasian population from central Russia. Molecular vision 21 34012228
2019 The protective variant rs7173049 at LOXL1 locus impacts on retinoic acid signaling pathway in pseudoexfoliation syndrome. Human molecular genetics 21 30986821
2013 Development of novel LOXL1 genotyping method and evaluation of LOXL1, APOE and MTHFR polymorphisms in exfoliation syndrome/glaucoma in a Greek population. Molecular vision 21 23687437
2011 Evaluation of LOXL1 polymorphisms in exfoliation syndrome in the Uygur population. Molecular vision 21 21738402
2021 lncRNA LOXL1-AS1 promotes liver cancer cell proliferation and migration by regulating the miR-377-3p/NFIB axis. Oncology letters 20 34267816
2017 Platelet-rich plasma induces post-natal maturation of immature articular cartilage and correlates with LOXL1 activation. Scientific reports 20 28623328
2016 Screening of lysyl oxidase (LOX) and lysyl oxidase like (LOXL) enzyme expression and activity in preterm prelabor rupture of fetal membranes. Journal of perinatal medicine 20 26011922
2009 From epidemiology to lysyl oxidase like one (LOXL1) polymorphisms discovery: phenotyping and genotyping exfoliation syndrome and exfoliation glaucoma in Iceland. Acta ophthalmologica 20 19664108
2022 Structural basis of lipoprotein recognition by the bacterial Lol trafficking chaperone LolA. Proceedings of the National Academy of Sciences of the United States of America 19 36037338
2021 LOXL1 gene polymorphisms are associated with exfoliation syndrome/exfoliation glaucoma risk: An updated meta-analysis. PloS one 19 33909695
2019 LOXL1-AS1 Drives The Progression Of Gastric Cancer Via Regulating miR-142-5p/PIK3CA Axis. OncoTargets and therapy 19 31908498
2014 Expression and regulation of LOXL1 and elastin-related genes in eyes with exfoliation syndrome. Journal of glaucoma 19 25275910
1989 Molecular genetics of human immune responsiveness to Lolium perenne (rye) allergen, Lol p III. International archives of allergy and applied immunology 19 2496037
2020 Identification and activity of the functional complex between hnRNPL and the pseudoexfoliation syndrome-associated lncRNA, LOXL1-AS1. Human molecular genetics 18 32037441
2020 Potential association of LOXL1 with peritoneal dissemination in gastric cancer possibly via promotion of EMT. PloS one 18 33095806
2019 LOXL1 folding in exfoliation glaucoma. Advances in protein chemistry and structural biology 18 31928728
2016 Baicalein Inhibits Epithelial to Mesenchymal Transition via Downregulation of Cyr61 and LOXL-2 in MDA-MB231 Breast Cancer Cells. Molecules and cells 18 28008161
2011 Review: The role of LOXL1 in exfoliation syndrome/glaucoma. Saudi journal of ophthalmology : official journal of the Saudi Ophthalmological Society 18 23960948
1996 Identification of T-cell epitopes of Lol p 9, a major allergen of ryegrass (Lolium perenne) pollen. The Journal of allergy and clinical immunology 18 8765826
1994 Mapping of T cell epitopes of the major fraction of rye grass using peripheral blood mononuclear cells from atopics and non-atopics. II. Isoallergen clone 5A of Lolium perenne group I (Lol p I). European journal of immunology 18 7522159
2021 LOXL1‑AS1 promotes thymoma and thymic carcinoma progression by regulating miR‑525‑5p‑HSPA9. Oncology reports 17 33907842

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