Affinage

PLOD2

Procollagen-lysine,2-oxoglutarate 5-dioxygenase 2 · UniProt O00469

Length
737 aa
Mass
84.7 kDa
Annotated
2026-06-10
100 papers in source corpus 23 papers cited in narrative 23 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 5/5 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

PLOD2 encodes telopeptide lysyl hydroxylase (LH2), a 2-oxoglutarate-dependent enzyme that hydroxylates lysine residues within the telopeptide domains of fibrillar collagens to enable formation of stable pyridinoline cross-links in the extracellular matrix (PMID:12881513). Loss-of-function mutations in PLOD2 cause Bruck syndrome, in which deficient telopeptide lysine hydroxylation produces a near-complete absence of stable trivalent collagen cross-links in bone with their replacement by allysine aldol dimers, while cartilage type II collagen is normally hydroxylated, revealing tissue-specific control of cross-link chemistry (PMID:12881513, PMID:33778323); a C-terminal hotspot (residues ~598-608) and the alternatively spliced exon 13a-containing isoform are specifically required for this catalytic function (PMID:15523624, PMID:22689593). PLOD2 is a hypoxia-responsive gene activated by HIF-1, and its activity drives ECM stiffening, collagen fiber alignment, and the downstream changes in tumor cell adhesion and motility that accompany matrix remodeling (PMID:23423382). Beyond collagen, PLOD2 hydroxylates integrin β1 to stabilize it and direct its plasma membrane localization, an event required for tumor cell motility and metastasis (PMID:32058962), and its enzymatic activity elevates cytoplasmic succinate that suppresses chromatin 5hmC to sustain mesenchymal and stemness phenotypes (PMID:37155842). In bone and mesenchymal progenitors, a HIF-1α/PLOD2/LOX axis couples collagen cross-linking to glycolytic metabolism and osteogenic differentiation (PMID:38472175).

Mechanistic history

Synthesis pass · year-by-year structured walk · 15 steps
  1. 2003 High

    Established the core molecular identity of PLOD2 as the telopeptide lysyl hydroxylase whose activity produces collagen pyridinoline cross-links, and tied its loss to human disease.

    Evidence Genetic mutation analysis of Bruck syndrome patients plus mRNA quantification and pyridinoline cross-link measurement in fibrotic fibroblasts

    PMID:12881513

    Open questions at the time
    • Did not resolve the structural basis of telopeptide specificity
    • Did not address tissue-specific differences in cross-link chemistry
  2. 2004 Medium

    Localized a C-terminal functional hotspot critical for hydroxylase activity and showed that alternative splicing of exon 13A generates isoforms determining cross-link type.

    Evidence Mutation mapping with urine cross-link metabolite analysis; RT-PCR/Northern blot with cycloheximide perturbation in fibroblasts and kidney cells

    PMID:15523624 PMID:15694128

    Open questions at the time
    • No enzyme assay directly linking the hotspot residues to catalysis
    • Factor controlling exon 13A inclusion not identified
  3. 2012 Medium

    Demonstrated that the exon 13a-containing long isoform is specifically required for normal bone telopeptide hydroxylation, refining the genotype-phenotype map.

    Evidence Homozygosity mapping and Sanger sequencing in consanguineous families

    PMID:22689593

    Open questions at the time
    • No direct enzyme assay of isoform-specific activity
    • Mechanism distinguishing Bruck syndrome from OI without contractures unclear
  4. 2013 High

    Placed PLOD2 in the hypoxia response by showing HIF-1 drives its expression to remodel and stiffen the ECM, connecting collagen biochemistry to tumor-relevant biomechanics.

    Evidence siRNA knockdown in fibroblasts with ECM stiffness/fiber alignment assays, cancer cell co-culture, and HIF-1 ChIP/reporter

    PMID:23423382

    Open questions at the time
    • Did not separate PLOD2 contribution from co-knocked P4HA1/P4HA2
    • Cross-link chemistry under hypoxia not directly measured
  5. 2016 Medium

    Identified post-transcriptional repression of PLOD2 by miR-26a/b, adding a layer of regulatory control over its tumor-promoting expression.

    Evidence Luciferase reporter on PLOD2 3'UTR plus miRNA restoration and migration/invasion assays in renal cell carcinoma

    PMID:26983694

    Open questions at the time
    • Single-lab luciferase validation
    • In vivo relevance of miR-26-PLOD2 axis not tested
  6. 2018 Medium

    Revealed a non-canonical nuclear-associated partnership of PLOD2 with the histone demethylase KDM1A regulating mineralization and odontogenic/osteogenic differentiation.

    Evidence Co-IP for KDM1A-PLOD2, siRNA knockdown, ALP/Alizarin red assays, and in vivo transplantation in stem cells of the apical papilla

    PMID:29656462

    Open questions at the time
    • Single Co-IP without reciprocal structural validation
    • Whether interaction depends on PLOD2 enzymatic activity unknown
  7. 2017 Medium

    Established multiple transcriptional inputs (HIF-1α, FOXA1, TGF-β1) converging on PLOD2 to drive EMT, migration, and invasion across cancer types.

    Evidence siRNA knockdown with EMT/PI3K-AKT marker Westerns, FOXA1 ChIP on the PLOD2 promoter, β-catenin/focal adhesion imaging, and in vivo metastasis models in glioma, NSCLC, and cervical cancer

    PMID:28410212 PMID:28507454 PMID:29072684

    Open questions at the time
    • EMT marker changes are correlative, not mechanistically dissected
    • Whether effects require collagen hydroxylation versus other substrates unresolved
  8. 2019 Medium

    Linked PLOD2 to cancer glucose metabolism through a STAT3-HK2 axis driving aerobic glycolysis.

    Evidence siRNA knockdown with HK2 overexpression rescue and glycolysis assays in colorectal cancer

    PMID:31742425

    Open questions at the time
    • Mechanism by which PLOD2 activates STAT3 not defined
    • Single lab
  9. 2020 High

    Defined a substrate beyond collagen: PLOD2 hydroxylates integrin β1 to stabilize and traffic it to the plasma membrane, mechanistically connecting PLOD2 enzymatic activity to cell motility and metastasis.

    Evidence Loss-of-function with subcellular fractionation, in vivo metastasis model, and patient tissue co-expression in head and neck SCC

    PMID:32058962

    Open questions at the time
    • Specific integrin β1 lysine sites not mapped
    • Generality across other adhesion receptors untested
  10. 2021 Medium

    Extended the integrin β1 axis upstream to the tumor microenvironment, showing M2-macrophage IL-6/STAT3 signaling drives PLOD2 to stabilize integrin β1, and characterized m6A-based post-transcriptional control via IGF2BP2.

    Evidence Luciferase reporter for STAT3 on PLOD2 promoter and tissue immunofluorescence in oral SCC; MeRIP/RIP-qPCR and CRISPR/dCas13b-ALKBH5 demethylation in varicocele testes

    PMID:33887877 PMID:37670228

    Open questions at the time
    • Single-lab promoter and reader validation
    • Cross-tissue generality of m6A control unclear
  11. 2021 Medium

    Showed PLOD2 acts as a required effector downstream of L1CAM/ezrin/SMAD2-3 in cancer and downstream of TGF-β1/Smad3 in tissue repair, demonstrating context-dependent placement in signaling cascades.

    Evidence shRNA/enzymatic inhibition with in vivo tumorigenesis and liver metastasis in colorectal cancer; mechanical injury and rat SCI models with TGF-β1 inhibitor and PLOD2 knockdown

    PMID:33495412 PMID:33805564

    Open questions at the time
    • Whether PLOD2 enzymatic substrate in these contexts is collagen or another protein unresolved
    • Single lab per context
  12. 2021 High

    Provided direct biochemical confirmation in patient tissue that PLOD2 loss specifically abolishes bone collagen telopeptide cross-links while sparing cartilage, defining tissue-specific cross-link chemistry.

    Evidence Mass spectrometry of bone collagen cross-linked peptides and urine-derived cartilage collagen from a Bruck syndrome patient

    PMID:33778323

    Open questions at the time
    • Molecular basis of cartilage-specific compensation not identified
    • Single patient
  13. 2023 High

    Uncovered an enzymatic-metabolic-epigenetic mechanism in which PLOD2 activity elevates cytoplasmic succinate to suppress chromatin 5hmC and sustain stemness, linking the dioxygenase to epigenetic reprogramming.

    Evidence Stable isotope metabolomics, 5hmC ChIP-seq, PLOD2 silencing, and exogenous succinate rescue in breast cancer

    PMID:37155842

    Open questions at the time
    • Enzymatic step generating excess succinate not pinpointed
    • Identity of 5hmC-writing dioxygenases inhibited not directly shown
  14. 2023 Medium

    Expanded PLOD2's protein-interaction repertoire to USP15 stabilization and EGFR binding driving AKT/mTOR signaling, and identified IRF5-PLOD2-BIRC3 as another regulatory axis, with minoxidil as a pharmacological inhibitor.

    Evidence Co-IP (USP15, EGFR), HIF1A/IRF5 promoter assays, AKT/mTOR phosphorylation Westerns, RNA-seq, in vivo xenografts, and minoxidil treatment in colorectal, renal, and hepatocellular carcinoma

    PMID:37227305 PMID:37577214 PMID:38008826

    Open questions at the time
    • Single Co-IP without reciprocal validation for each interaction
    • Whether these interactions require PLOD2 catalytic activity unknown
  15. 2024 High

    Integrated PLOD2 into a HIF-1α/PLOD2/LOX axis in mesenchymal progenitors coupling collagen cross-linking to glycolysis and osteogenic differentiation, and defined HIF-1α-driven METTL3-dependent m6A translational control of PLOD2.

    Evidence Conditional Hif1α knockout mouse with scRNA-seq and PLOD2 inhibition in heterotopic ossification; MeRIP-seq, METTL3 inhibitor, and AAV9-PLOD2 rescue in renal cell carcinoma

    PMID:38233403 PMID:38472175

    Open questions at the time
    • Direct enzymatic link between PLOD2 and glycolytic flux not fully defined
    • How LOX cooperates mechanistically with PLOD2 unresolved

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved which PLOD2 substrate (collagen telopeptide lysines versus integrin β1 versus other proteins) and which activity (hydroxylation versus succinate generation versus protein scaffolding) dominates in each disease and cancer context.
  • No structure of human PLOD2 with substrate
  • Substrate site-resolution for non-collagen targets missing
  • Catalytic dependence of reported protein-protein interactions untested

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0016491 oxidoreductase activity 3 GO:0016787 hydrolase activity 1 GO:0140096 catalytic activity, acting on a protein 1
Localization
GO:0005576 extracellular region 2 GO:0005829 cytosol 2
Pathway
R-HSA-1474244 Extracellular matrix organization 3 R-HSA-1643685 Disease 3
Partners
EGFRIGF2BP2ITGB1KDM1AUSP15

Evidence

Reading pass · 23 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2003 PLOD2 was identified as the telopeptide lysyl hydroxylase (TLH) responsible for hydroxylation of lysine residues within the telopeptides of fibrillar collagens. Two missense mutations in exon 17 of PLOD2 were found in Bruck syndrome patients who have pyridinoline cross-link deficiency in bone collagen, and fibrotic fibroblasts with elevated PLOD2 mRNA showed increased pyridinoline cross-link levels in deposited matrix, directly linking PLOD2 enzymatic activity to telopeptide lysine hydroxylation and collagen cross-linking. Genetic mutation analysis of Bruck syndrome patients; mRNA quantification and pyridinoline cross-link measurement in SSc fibroblasts The Journal of biological chemistry High 12881513
2004 A hotspot region in the C-terminal domain of PLOD2 (residues ~598–608) is functionally critical for telopeptide lysyl hydroxylase activity; missense mutations (Arg598His, Gly601Val, Thr608Ile) in this region cause Bruck syndrome with pyridinoline deficiency and low collagen cross-link degradation products in urine. Mutation analysis of Bruck syndrome patient with Arg598His substitution; urine collagen cross-link metabolite analysis American journal of medical genetics. Part A Medium 15523624
2004 PLOD2 exists as two alternatively spliced isoforms: a long form (including 63 bp exon 13A, ubiquitously expressed) and a short form (co-expressed in kidney, spleen, liver, placenta). The splicing of exon 13A is regulated by cell density and protein synthesis (cycloheximide treatment), suggesting a newly-synthesized factor controls inclusion of exon 13A. The relative expression of these isoforms determines the type of collagen cross-links formed. RT-PCR, Northern blot; cycloheximide treatment of human skin fibroblasts and kidney cells Matrix biology : journal of the International Society for Matrix Biology Medium 15694128
2012 Homozygous frameshift mutations in the alternative exon 13a of PLOD2 are sufficient to cause Bruck syndrome, indicating that the longer PLOD2 protein isoform (encoded by exon 13a) is specifically required for normal telopeptide lysine hydroxylation in bone. Compound heterozygous PLOD2 mutations can also produce autosomal recessive OI phenotypes without joint contractures. Homozygosity mapping, Sanger sequencing, mutation analysis in consanguineous families Human mutation Medium 22689593
2013 HIF-1 transcriptionally activates PLOD2 expression in fibroblasts under hypoxic conditions. PLOD2 is required for ECM stiffening and collagen fiber alignment under hypoxia; its loss (along with P4HA1/P4HA2) prevents hypoxia-induced ECM remodeling, collagen deposition, and downstream changes in breast cancer cell morphology, adhesion, and motility. siRNA knockdown of PLOD2 in fibroblasts; collagen fiber alignment and ECM stiffness assays; co-culture with breast cancer cells; HIF-1 ChIP/reporter assays The Journal of biological chemistry High 23423382
2021 Compound heterozygous PLOD2 mutations in a Bruck syndrome patient cause diminished hydroxylation specifically of type I collagen telopeptide lysines (but not triple-helical lysines), resulting in near-complete absence of stable trivalent collagen cross-links in bone and replacement by allysine aldol dimeric cross-links (as in skin). Cartilage type II collagen telopeptide lysines from the same patient were normally hydroxylated, revealing tissue-specific control of collagen cross-link chemistry. Mass spectrometry of bone collagen cross-linked peptides from patient biopsy; comparison with urine-derived cartilage collagen JBMR plus High 33778323
2017 PLOD2 knockdown in glioma cells inactivates the PI3K/AKT signaling pathway and downregulates EMT-associated markers (E-cadherin, vimentin, N-cadherin, β-catenin, snail, slug), inhibiting proliferation, migration, and invasion. PLOD2 expression is induced by HIF-1α under hypoxia, placing PLOD2 downstream of HIF-1α in hypoxia-driven EMT. siRNA knockdown; Western blot for PI3K/AKT and EMT markers; in vitro migration/invasion assays; in vivo xenograft Oncotarget Medium 28410212
2017 PLOD2 is transcriptionally regulated by the transcription factor FOXA1 acting downstream of the EGFR-PI3K/AKT axis. FOXA1 directly binds the PLOD2 promoter to activate PLOD2 transcription in NSCLC cells. PLOD2 promotes metastasis directly by enhancing migration and indirectly by inducing collagen reorganization. ChIP assay of FOXA1 on PLOD2 promoter; gain- and loss-of-function studies; orthotopic metastasis model Cell death & disease Medium 29072684
2017 PLOD2 knockdown in cervical cancer cells inhibits cell migration, invasion, and adhesion to type I collagen. PLOD2 depletion prevents β-catenin nuclear translocation (blocking TGF-β1-induced EMT) and inhibits focal adhesion formation. Hypoxia and TGF-β1 each induce PLOD2 expression to promote these processes. siRNA knockdown; wound healing/Transwell assays; immunocytochemistry for β-catenin; phalloidin staining for focal adhesions; Western blot Cancer cell international Medium 28507454
2020 PLOD2 hydroxylates integrin β1, and this hydroxylation is required for integrin β1 stabilization and recruitment to the plasma membrane. Loss of PLOD2 expression prevents integrin β1 from reaching its functional site (plasma membrane), abrogating tumor cell motility and metastasis in vivo in head and neck SCC. Loss-of-function studies; subcellular fractionation/membrane localization of integrin β1; in vivo metastasis model; co-expression analysis in patient tissue iScience High 32058962
2021 IL-6 released by CD163+ M2-type tumor-associated macrophages activates STAT3, which directly binds and activates the PLOD2 promoter in oral SCC cells, thereby increasing PLOD2 expression. This upregulation of PLOD2 stabilizes integrin β1 and promotes invasion/metastasis. IL-6 stimulation did not directly affect integrin β1 expression or maturation. Dual-luciferase reporter assay for STAT3 binding to PLOD2 promoter; siRNA; immunofluorescence double staining for IL-6 and CD163 in patient tissue International journal of oncology Medium 33887877
2023 PLOD2 expression is associated with elevated cytoplasmic succinate levels during EMT. PLOD2 silencing in breast cancer cells reduces succinate levels and inhibits mesenchymal phenotypes and stemness, accompanied by increased 5-hydroxymethylcytosine (5hmC) in chromatin. Exogenous succinate rescues stemness and 5hmC levels in PLOD2-silenced cells, indicating PLOD2 promotes cancer progression at least partly through succinate-mediated epigenetic regulation. Stable isotope-resolved metabolomics; ChIP-seq for 5hmC; PLOD2 silencing; exogenous succinate rescue experiment Proceedings of the National Academy of Sciences of the United States of America High 37155842
2023 PLOD2 interacts with USP15 (ubiquitin-specific protease 15) and stabilizes USP15 in the cytoplasm, leading to activation of AKT/mTOR phosphorylation and promotion of colorectal cancer proliferation, invasion, and metastasis. The PLOD2 inhibitor minoxidil downregulates PLOD2, suppresses USP15, and inhibits AKT/mTOR phosphorylation. Co-immunoprecipitation; Western blot for AKT/mTOR phosphorylation; overexpression and knockdown; in vivo xenograft; minoxidil pharmacological inhibition Cancer science Medium 37227305
2023 PLOD2 binds to the extracellular domain of EGFR, leading to EGFR phosphorylation and downstream AKT signaling pathway activation in clear cell renal cell carcinoma. HIF1A transcriptionally activates PLOD2 by binding a specific promoter region. PLOD2 inhibitor minoxidil suppresses ccRCC progression by inactivating the EGFR/AKT axis. Co-immunoprecipitation (PLOD2–EGFR interaction); HIF1A ChIP on PLOD2 promoter; phosphorylation assays; knockdown/overexpression; in vivo xenograft; minoxidil treatment Cell death & disease Medium 38008826
2019 PLOD2 promotes aerobic glycolysis in colorectal cancer cells by upregulating hexokinase 2 (HK2) expression via the STAT3 signaling pathway. HK2 overexpression reverses the inhibitory effects of PLOD2 knockdown, establishing a PLOD2–STAT3–HK2 functional axis in glucose metabolism. siRNA knockdown of PLOD2; HK2 overexpression rescue; Western blot for STAT3 and HK2; glycolysis assays Biochemistry and cell biology Medium 31742425
2018 KDM1A (histone demethylase) associates with PLOD2 to form a protein complex in stem cells of the apical papilla (SCAPs). Knockdown of either KDM1A or PLOD2 inhibits alkaline phosphatase activity and mineralization and promotes expression of osteo/dentinogenic differentiation markers (DSPP, DMP1, BSP, RUNX2, OSX, DLX2). Co-immunoprecipitation (KDM1A–PLOD2 interaction); siRNA knockdown; ALP assay; Alizarin red staining; in vivo transplantation Cell proliferation Medium 29656462
2024 METTL3-mediated m6A modification of PLOD2 mRNA at its 3'UTR promotes PLOD2 protein translation under prolonged hypoxia. HIF-1α binds the METTL3 promoter to enhance its transcription, and METTL3 in turn increases PLOD2 protein levels in an m6A-dependent manner to promote renal cell carcinoma progression. AAV9-mediated re-transduction of PLOD2 rescued tumor growth suppressed by METTL3 inhibition in vivo. MeRIP-seq; METTL3 inhibitor STM2457; AAV9-PLOD2 rescue in vivo; HIF-1α ChIP on METTL3 promoter; Western blot Cell death & disease Medium 38233403
2021 IGF2BP2 binds to the m6A-modified 3'UTR of PLOD2 mRNA and positively regulates PLOD2 mRNA stability in varicocele rat testes. Targeted demethylation of PLOD2 m6A by a CRISPR/dCas13b-ALKBH5 system decreases PLOD2 expression and promotes GC-2 cell proliferation while inhibiting apoptosis under oxidative stress. MeRIP-seq/qPCR; RIP-qPCR; luciferase assay for IGF2BP2–PLOD2 mRNA interaction; CRISPR/dCas13b-ALKBH5 demethylation; cell proliferation/apoptosis assays Cellular & molecular biology letters Medium 37670228
2021 PLOD2 is required for L1CAM-driven colorectal cancer proliferation, motility, tumorigenesis, and liver metastasis. L1CAM upregulates PLOD2 expression via ezrin signaling and SMAD2/3 pathway activation. Suppression of endogenous PLOD2 expression or inhibition of its enzymatic activity blocks the tumorigenic properties conferred by L1CAM. shRNA-mediated PLOD2 knockdown; enzymatic inhibition; tumorigenesis/liver metastasis assays in vivo; Western blot for SMAD2/3 International journal of molecular sciences Medium 33805564
2021 Adipose-derived stromal cells (ADSCs) upregulate PLOD2 in spinal cord neurons via TGF-β1 secretion and downstream Smad3 phosphorylation. PLOD2 upregulation is required for the neuroprotective/neurorestorative effects of ADSCs after spinal cord injury; inhibiting TGF-β1 signaling (with SB431542) or PLOD2 expression abrogates neuronal survival, axonal regeneration promotion, and functional recovery. In vitro mechanical injury model; rat SCI model; PLOD2 knockdown; TGF-β1 receptor inhibitor SB431542; immunostaining for MAP2, NSE, GAP43, GFAP Aging Medium 33495412
2024 HIF-1α activates PLOD2 expression in mesenchymal progenitor cells, and the HIF-1α/PLOD2/LOX axis regulates collagen cross-linking, glucose metabolism, and osteogenic differentiation. Inhibition of PLOD2 activity significantly decreases osteogenic differentiation and glycolytic metabolism in MPCs, and genetic deletion of Hif1α in zeugopod mesenchymal cells reduces PLOD2/LOX pathway activity and mitigates heterotopic ossification in vivo. Conditional Hif1α knockout mouse (Hoxa11-CreERT2; Hif1afl/fl); scRNA-seq; pharmacological PLOD2 inhibition; osteogenic differentiation assays; glycolysis measurement Bone research High 38472175
2023 PLOD2 is transcriptionally regulated by interferon regulatory factor 5 (IRF5) in hepatocellular carcinoma cells, as shown by a luciferase reporter assay. PLOD2 in turn promotes HCC metastasis via upregulation of BIRC3 (baculoviral IAP repeat containing 3), placing PLOD2 in an IRF5–PLOD2–BIRC3 signaling axis. Luciferase reporter assay (IRF5 → PLOD2 promoter); RNA sequencing; PLOD2 knockdown with downstream BIRC3 measurement; in vivo metastasis model Journal of clinical and translational hepatology Medium 37577214
2016 miR-26a-5p and miR-26b-5p directly target the PLOD2 3'UTR to suppress its expression, as validated by luciferase reporter assay. Downregulation of PLOD2 by these miRNAs significantly inhibits cancer cell migration and invasion in renal cell carcinoma. In silico analysis; luciferase reporter assay; miRNA restoration in RCC cell lines; migration/invasion assays; PLOD2 siRNA knockdown International journal of oncology Medium 26983694

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2013 Hypoxia-inducible factor 1 (HIF-1) promotes extracellular matrix remodeling under hypoxic conditions by inducing P4HA1, P4HA2, and PLOD2 expression in fibroblasts. The Journal of biological chemistry 394 23423382
2003 Identification of PLOD2 as telopeptide lysyl hydroxylase, an important enzyme in fibrosis. The Journal of biological chemistry 319 12881513
1993 LH-2: a LIM/homeodomain gene expressed in developing lymphocytes and neural cells. Proceedings of the National Academy of Sciences of the United States of America 183 7678338
2016 Tumour-suppressive miRNA-26a-5p and miR-26b-5p inhibit cell aggressiveness by regulating PLOD2 in bladder cancer. British journal of cancer 132 27310702
2004 Phenotypic and molecular characterization of Bruck syndrome (osteogenesis imperfecta with contractures of the large joints) caused by a recessive mutation in PLOD2. American journal of medical genetics. Part A 132 15523624
1992 Mutants of Rhodobacter sphaeroides lacking one or more pigment-protein complexes and complementation with reaction-centre, LH1, and LH2 genes. Molecular microbiology 131 1588816
1997 Apoprotein structure in the LH2 complex from Rhodopseudomonas acidophila strain 10050: modular assembly and protein pigment interactions. Journal of molecular biology 130 9159480
2012 Quantum coherence spectroscopy reveals complex dynamics in bacterial light-harvesting complex 2 (LH2). Proceedings of the National Academy of Sciences of the United States of America 122 22215585
2000 Influence of anionic ligands (X) on the nature and magnetic properties of dinuclear LCuDgX3.nH2O complexes (LH2 standing for tetradentate Schiff base ligands deriving from 2-hydroxy-3-methoxybenzaldehyde and X being Cl, N3C2, and CF3COO). Inorganic chemistry 120 11272520
2013 High frequency of virulence factor genes tdh, trh, and tlh in Vibrio parahaemolyticus strains isolated from a pristine estuary. Applied and environmental microbiology 115 23354697
1998 Expression of the LIM-homeobox gene LH2 generates immortalized steel factor-dependent multipotent hematopoietic precursors. The EMBO journal 113 9755174
2004 Variable LH2 stoichiometry and core clustering in native membranes of Rhodospirillum photometricum. The EMBO journal 100 15457213
1994 The relationship between carotenoid biosynthesis and the assembly of the light-harvesting LH2 complex in Rhodobacter sphaeroides. The Biochemical journal 94 8129720
2017 Hypoxia-induced PLOD2 promotes proliferation, migration and invasion via PI3K/Akt signaling in glioma. Oncotarget 86 28410212
2017 PLOD2 in cancer research. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 82 28415047
2017 PLOD2 regulated by transcription factor FOXA1 promotes metastasis in NSCLC. Cell death & disease 82 29072684
2017 Hypoxia and TGF-β1 induced PLOD2 expression improve the migration and invasion of cervical cancer cells by promoting epithelial-to-mesenchymal transition (EMT) and focal adhesion formation. Cancer cell international 74 28507454
2016 Regulation of the collagen cross-linking enzymes LOXL2 and PLOD2 by tumor-suppressive microRNA-26a/b in renal cell carcinoma. International journal of oncology 74 26983694
2012 Mutations in PLOD2 cause autosomal-recessive connective tissue disorders within the Bruck syndrome--osteogenesis imperfecta phenotypic spectrum. Human mutation 72 22689593
1999 Evaluation of alkaline phosphatase- and digoxigenin-labelled probes for detection of the thermolabile hemolysin (tlh) gene of Vibrio parahaemolyticus. Letters in applied microbiology 70 10030035
2015 Antioxidant and anti-aging activities of the polysaccharide TLH-3 from Tricholoma lobayense. International journal of biological macromolecules 59 26721384
2003 AFM characterization of tilt and intrinsic flexibility of Rhodobacter sphaeroides light harvesting complex 2 (LH2). Journal of molecular biology 57 12498803
2016 An Ab Initio Description of the Excitonic Properties of LH2 and Their Temperature Dependence. The journal of physical chemistry. B 56 27791372
1997 Influence of the protein binding site on the absorption properties of the monomeric bacteriochlorophyll in Rhodobacter sphaeroides LH2 complex. Biochemistry 52 9405063
2017 Immunomodulating activity of the polysaccharide TLH-3 from Tricholomalobayense in RAW264.7 macrophages. International journal of biological macromolecules 48 29155198
2003 Detergent structure in crystals of the integral membrane light-harvesting complex LH2 from Rhodopseudomonas acidophila strain 10050. Journal of molecular biology 48 12547211
2013 Genes similar to the Vibrio parahaemolyticus virulence-related genes tdh, tlh, and vscC2 occur in other vibrionaceae species isolated from a pristine estuary. Applied and environmental microbiology 43 24212573
2014 Static and Dynamic Disorder in Bacterial Light-Harvesting Complex LH2: A 2DES Simulation Study. The journal of physical chemistry. B 41 24945779
2025 Single-cell RNA sequencing and immune microenvironment analysis reveal PLOD2-driven malignant transformation in cervical cancer. Frontiers in immunology 40 39840054
2000 Three different turkey luteinizing hormone receptor (tLH-R) isoforms I: characterization of alternatively spliced tLH-R isoforms and their regulated expression in diverse tissues. Biology of reproduction 39 10611074
2018 Engineering of B800 bacteriochlorophyll binding site specificity in the Rhodobacter sphaeroides LH2 antenna. Biochimica et biophysica acta. Bioenergetics 38 30414933
2007 Directed formation of micro- and nanoscale patterns of functional light-harvesting LH2 complexes. Journal of the American Chemical Society 37 17985885
2006 Theoretical prediction of spectral and optical properties of bacteriochlorophylls in thermally disordered LH2 antenna complexes. The Journal of chemical physics 37 16863329
2004 Tissue-specific expression and regulation of the alternatively-spliced forms of lysyl hydroxylase 2 (LH2) in human kidney cells and skin fibroblasts. Matrix biology : journal of the International Society for Matrix Biology 37 15694128
2018 PLOD2 as a potential regulator of peritoneal dissemination in gastric cancer. International journal of cancer 35 29603227
2020 PLOD2 Is Essential to Functional Activation of Integrin β1 for Invasion/Metastasis in Head and Neck Squamous Cell Carcinomas. iScience 34 32058962
2019 PLOD2 promotes aerobic glycolysis and cell progression in colorectal cancer by upregulating HK2. Biochemistry and cell biology = Biochimie et biologie cellulaire 34 31742425
2023 The PLOD2/succinate axis regulates the epithelial-mesenchymal plasticity and cancer cell stemness. Proceedings of the National Academy of Sciences of the United States of America 33 37155842
2018 Hypoxia-Induced PLOD2 is a Key Regulator in Epithelial-Mesenchymal Transition and Chemoresistance in Biliary Tract Cancer. Annals of surgical oncology 33 30105440
2023 Apelin promotes osteosarcoma metastasis by upregulating PLOD2 expression via the Hippo signaling pathway and hsa_circ_0000004/miR-1303 axis. International journal of biological sciences 32 36632453
2012 Electrochemical deoxyribonucleic acid biosensor based on carboxyl functionalized graphene oxide and poly-L-lysine modified electrode for the detection of tlh gene sequence related to vibrio parahaemolyticus. Analytica chimica acta 30 23101650
2002 Dynamics of energy transfer from lycopene to bacteriochlorophyll in genetically-modified LH2 complexes of Rhodobacter sphaeroides. Biochemistry 30 11900556
2023 PLOD2 promotes colorectal cancer progression by stabilizing USP15 to activate the AKT/mTOR signaling pathway. Cancer science 29 37227305
2003 Membrane protein stability: high pressure effects on the structure and chromophore-binding properties of the light-harvesting complex LH2. Biochemistry 29 14596617
2008 Static and dynamic protein impact on electronic properties of light-harvesting complex LH2. The journal of physical chemistry. B 28 19367872
2004 In vitro self-assembly of the light harvesting pigment-protein LH2 revealed by ultrafast spectroscopy and electron microscopy. Biophysical journal 28 15041674
2021 PLOD2-driven IL-6/STAT3 signaling promotes the invasion and metastasis of oral squamous cell carcinoma via activation of integrin β1. International journal of oncology 27 33887877
2013 The LH2 complexes are assembled in the cells of purple sulfur bacterium Ectothiorhodospira haloalkaliphila with inhibition of carotenoid biosynthesis. Photosynthesis research 27 24163008
2007 Comparison of the fluorescence kinetics of detergent-solubilized and membrane-reconstituted LH2 complexes from Rps. acidophila and Rb. sphaeroides. Photosynthesis research 27 17912609
2023 Hypoxia-induced PLOD2 promotes clear cell renal cell carcinoma progression via modulating EGFR-dependent AKT pathway activation. Cell death & disease 26 38008826
2021 Abnormal Bone Collagen Cross-Linking in Osteogenesis Imperfecta/Bruck Syndrome Caused by Compound Heterozygous PLOD2 Mutations. JBMR plus 26 33778323
2019 PLOD2 contributes to drug resistance in laryngeal cancer by promoting cancer stem cell-like characteristics. BMC cancer 26 31455288
2017 Association of ECRG4 with PLK1, CDK4, PLOD1 and PLOD2 in esophageal squamous cell carcinoma. American journal of translational research 25 28861165
2024 The HIF-1α/PLOD2 axis integrates extracellular matrix organization and cell metabolism leading to aberrant musculoskeletal repair. Bone research 24 38472175
2019 Use of Mass Spectrometry to Profile Peptides in Whey Protein Isolate Medium Fermented by Lactobacillus helveticus LH-2 and Lactobacillus acidophilus La-5. Frontiers in nutrition 24 31681785
2011 Monomeric RC-LH1 core complexes retard LH2 assembly and intracytoplasmic membrane formation in PufX-minus mutants of Rhodobacter sphaeroides. Biochimica et biophysica acta 24 21663730
2019 Hypoxia-induced PLOD2 regulates invasion and epithelial-mesenchymal transition in endometrial carcinoma cells. Genes & genomics 23 31872384
2011 A comparative spectroscopic and kinetic study of photoexcitations in detergent-isolated and membrane-embedded LH2 light-harvesting complexes. Biochimica et biophysica acta 23 22172735
2018 KDM1A regulated the osteo/dentinogenic differentiation process of the stem cells of the apical papilla via binding with PLOD2. Cell proliferation 22 29656462
2015 Functional characteristics of spirilloxanthin and keto-bearing Analogues in light-harvesting LH2 complexes from Rhodobacter sphaeroides with a genetically modified carotenoid synthesis pathway. Biochimica et biophysica acta 22 25871644
2014 The origin of the split B800 absorption peak in the LH2 complexes from Allochromatium vinosum. Photosynthesis research 22 25150556
2009 Spectral diffusion and electron-phonon coupling of the B800 BChl a molecules in LH2 complexes from three different species of purple bacteria. Biophysical journal 22 19883604
2022 circRNA PLOD2 promotes tumorigenesis and Warburg effect in colon cancer by the miR-513a-5p/SIX1/LDHA axis. Cell cycle (Georgetown, Tex.) 21 36071678
2021 The Collagen-Modifying Enzyme PLOD2 Is Induced and Required during L1-Mediated Colon Cancer Progression. International journal of molecular sciences 21 33805564
2024 METTL3 facilitates renal cell carcinoma progression by PLOD2 m6A-methylation under prolonged hypoxia. Cell death & disease 20 38233403
2024 Development of Multienzyme Isothermal Rapid Amplification (MIRA) Combined with Lateral-Flow Dipstick (LFD) Assay to Detect Species-Specific tlh and Pathogenic trh and tdh Genes of Vibrio parahaemolyticus. Pathogens (Basel, Switzerland) 20 38251364
2021 Adipose-derived stromal cells improve functional recovery after spinal cord injury through TGF-β1/Smad3/PLOD2 pathway activation. Aging 20 33495412
2017 Novel Mutations in PLOD2 Cause Rare Bruck Syndrome. Calcified tissue international 20 29177700
2008 Femtosecond spectroscopy of native and carotenoidless purple-bacterial LH2 clarifies functions of carotenoids. Biophysical journal 20 18339744
2022 Tensor Network Path Integral Study of Dynamics in B850 LH2 Ring with Atomistically Derived Vibrations. Journal of chemical theory and computation 19 35732015
2019 Bruck syndrome 2 variant lacking congenital contractures and involving a novel compound heterozygous PLOD2 mutation. Bone 19 31472299
2017 Intermolecular Modes between LH2 Bacteriochlorophylls and Protein Residues: The Effect on the Excitation Energies. The journal of physical chemistry. B 18 28485594
2007 Solvation effect of bacteriochlorophyll excitons in light-harvesting complex LH2. Biophysical journal 18 17513366
2021 Intraband dynamics and exciton trapping in the LH2 complex of Rhodopseudomonas acidophila. The Journal of chemical physics 16 33514092
2021 Targeted Demethylation of the PLOD2 mRNA Inhibits the Proliferation and Migration of Renal Cell Carcinoma. Frontiers in molecular biosciences 16 34179084
2018 A Novel qPCR Method for Simultaneous Detection and Quantification of Viable Pathogenic and Non-pathogenic Vibrio parahaemolyticus (tlh , tdh , and ureR ). Frontiers in microbiology 16 30116230
2018 DT-13 suppresses breast cancer metastasis by modulating PLOD2 in the adipocytes microenvironment. Phytomedicine : international journal of phytotherapy and phytopharmacology 16 31005809
2015 Antioxidant capacity and cytotoxicity of sulfated polysaccharide TLH-3 from Tricholoma lobayense. International journal of biological macromolecules 16 26449529
2015 Fluorescence polarization measures energy funneling in single light-harvesting antennas--LH2 vs conjugated polymers. Scientific reports 16 26478272
2013 Fluorescence-excitation and emission spectra from LH2 antenna complexes of Rhodopseudomonas acidophila as a function of the sample preparation conditions. The journal of physical chemistry. B 16 24033126
2012 Population and coherence dynamics in light harvesting complex II (LH2). The Journal of chemical physics 16 22938221
2010 Expression, purification, and characterization of thermolabile hemolysin (TLH) from Vibrio alginolyticus. Diseases of aquatic organisms 16 20662368
2021 Virulence factor genes and comparative pathogenicity study of tdh, trh and tlh positive Vibrio parahaemolyticus strains isolated from Whiteleg shrimp, Litopenaeus vannamei (Boone, 1931) in India. Infection, genetics and evolution : journal of molecular epidemiology and evolutionary genetics in infectious diseases 15 34536578
2018 Adaptation of Rhodopseudomonas acidophila strain 7050 to growth at different light intensities: what are the benefits to changing the type of LH2? Faraday discussions 15 29355274
2008 Protein-induced geometric constraints and charge transfer in bacteriochlorophyll-histidine complexes in LH2. Physical chemistry chemical physics : PCCP 15 19030592
2001 Probing the binding sites of exchanged chlorophyll a in LH2 by Raman and site-selection fluorescence spectroscopies. FEBS letters 15 11226437
1989 Characterization and complementation of a mutant of Rhodobacter sphaeroides with a chromosomal deletion in the light-harvesting (LH2) genes. Journal of general microbiology 15 2693605
2023 Spindle cell rhabdomyosarcomas: With TFCP2 rearrangements, and novel EWSR1::ZBTB41 and PLOD2::RBM6 gene fusions. A study of five cases and review of the literature. Histopathology 14 38114270
2022 Pan-Cancer Analyses Reveal Oncogenic and Immunological Role of PLOD2. Frontiers in genetics 14 35586565
2022 Silencing PLOD2 attenuates cancer stem cell-like characteristics and cisplatin-resistant through Integrin β1 in laryngeal cancer. Translational oncology 14 35660870
2021 CircRNA PLOD2 enhances ovarian cancer propagation by controlling miR-378. Saudi journal of biological sciences 14 34759745
2020 PLOD2 increases resistance of gastric cancer cells to 5-fluorouracil by upregulating BCRP and inhibiting apoptosis. Journal of Cancer 14 32284742
2023 tDCS Regulates ASBT-3-OxoLCA-PLOD2-PTEN Signaling Pathway to Confer Neuroprotection Following Rat Cerebral Ischemia-Reperfusion Injury. Molecular neurobiology 13 37477767
2023 Dysregulation of PLOD2 Promotes Tumor Metastasis and Invasion in Hepatocellular Carcinoma. Journal of clinical and translational hepatology 13 37577214
2009 The telomere-linked helicase (TLH) gene family in Magnaporthe oryzae: revised gene structure reveals a novel TLH-specific protein motif. Current genetics 13 19360408
2023 N6-methyladenosine modification of PLOD2 causes spermatocyte damage in rats with varicocele. Cellular & molecular biology letters 12 37670228
2006 Carotenoid stoichiometry in the LH2 crystal: no spectral evidence for the presence of the second molecule in the alpha/beta-apoprotein dimer. FEBS letters 12 16790242
2001 Self diffusion and spectral modifications of a membrane protein, the Rubrivivax gelatinosus LH2 complex, incorporated into a monoolein cubic phase. Biophysical journal 12 11509374
2017 Emended description of the family Chromatiaceae, phylogenetic analyses of the genera Alishewanella, Rheinheimera and Arsukibacterium, transfer of Rheinheimera longhuensis LH2-2T to the genus Alishewanella and description of Alishewanella alkalitolerans sp. nov. from Lonar Lake, India. Antonie van Leeuwenhoek 11 28612170
2005 Temperature broadening of LH2 absorption in glycerol solution. Photosynthesis research 11 16172925

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