Affinage

MT2A

Metallothionein-2 · UniProt P02795

Length
61 aa
Mass
6.0 kDa
Annotated
2026-06-10
40 papers in source corpus 16 papers cited in narrative 16 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

MT2A is a cysteine-rich metal-binding protein that buffers intracellular zinc and copper and, through these metal pools and a set of protein partners, governs cell-fate decisions of survival, apoptosis, differentiation, and metabolism (PMID:28507149, PMID:17224279). The purified protein intrinsically binds metal, scavenges hydroxyl radicals, and protects DNA against UVC damage, establishing its core cytoprotective biochemistry (PMID:17224279). Its free-zinc-buffering role is set by competing partners: XAF1 binds MT2A and routes it to lysosomal degradation, raising free zinc, activating p53, and inactivating XIAP to drive apoptosis, while MT2A reciprocally destabilizes XAF1, creating a mutual antagonism that titrates the stress response (PMID:28507149); HMBOX1 binding instead elevates free zinc to support anti-apoptotic and pro-autophagic outcomes in endothelial cells (PMID:26456220). MT2A also acts in copper handling and cuproptosis: it limits intracellular Cu2+ and suppresses cuproptosis markers (FDX1, CTR1, DLAT), with Lys-31 identified as a functional site and direct small-molecule (artesunate) binding demonstrated (PMID:40754045), and it protects mitochondria from copper-overload toxicity in endothelial and ischemic-angiogenesis models (PMID:39915841). Under hypoxia MT2A translocates to mitochondria in a copper-dependent manner and binds tetrameric PKM2 to sustain its activity and promote glycolysis and OXPHOS (PMID:41211480). Through partner and pathway control, MT2A engages BARD1/BRCA1 in chemoresistance (PMID:32638210), the MST1/LATS2/YAP1 Hippo axis (PMID:35642057), NF-κB (PMID:34220318), and p38 MAPK signaling (PMID:35643179, PMID:38546949), the latter underlying neuronal anti-apoptotic protection (PMID:38546949). MT2A transcription is controlled by an HSF1 promoter-binding activator that is itself destabilized by the RING1 E3 ligase (PMID:37797799) and by lncRNA-directed recruitment of EZH2/LSD1 chromatin-modifying activity to its promoter (PMID:37548351).

Mechanistic history

Synthesis pass · year-by-year structured walk · 10 steps
  1. 2006 Medium

    Established that MT2A's protective activities are intrinsic to the protein, defining its core biochemistry of metal binding, radical scavenging, and DNA protection rather than dependence on cellular context.

    Evidence In vitro metal-binding, hydroxyl radical scavenging, and UVC DNA-damage protection assays with purified recombinant human protein

    PMID:17224279

    Open questions at the time
    • Single-lab in vitro study
    • Does not address which metals dominate in vivo or stoichiometry under physiological conditions
  2. 2014 Medium

    Linked MT2A to inflammatory signaling by identifying EOLA1 as an interactor and showing MT2A is required for LPS-induced IL-6 in endothelial cells.

    Evidence Yeast two-hybrid screen plus EOLA1/MT2A knockdown with IL-6 and apoptosis readouts in HUVECs

    PMID:24916366

    Open questions at the time
    • Interaction not confirmed by Co-IP in this study
    • Molecular mechanism connecting MT2A to IL-6 transcription not defined
  3. 2015 Medium

    Showed that partner identity tunes MT2A's free-zinc output toward survival, with HMBOX1 binding raising zinc to drive anti-apoptotic and pro-autophagic effects, and extended the EOLA1 axis to VCAM-1.

    Evidence Yeast two-hybrid, reciprocal Co-IP, MT2A knockdown with zinc chelation rescue (HMBOX1); knockdown/overexpression with VCAM-1 readout (EOLA1) in endothelial cells

    PMID:26456220 PMID:26881174

    Open questions at the time
    • VCAM-1 finding is Low confidence with no biochemical interaction validation
    • Structural basis of HMBOX1-driven zinc elevation unknown
  4. 2017 High

    Defined the mechanism by which MT2A-controlled zinc dictates apoptosis versus survival, showing XAF1 degrades MT2A to release zinc and activate p53/XIAP, with reciprocal MT2A destabilization of XAF1.

    Evidence Co-IP, binding-deficient XAF1 mutant, lysosomal degradation and zinc-measurement assays, and xenograft loss-of-function models

    PMID:28507149

    Open questions at the time
    • Does not resolve how cells choose between XAF1-driven degradation and HMBOX1-driven stabilization
    • Lysosomal targeting machinery for MT2A not identified
  5. 2019 Low

    Proposed epigenetic silencing of MT2A via lncRNA-directed EZH2 recruitment to its promoter.

    Evidence RIP, ChIP for EZH2 on the MT2A promoter, and TUG1 knockdown (paper subsequently retracted)

    PMID:30787623

    Open questions at the time
    • Paper was retracted, so the finding cannot be relied upon
    • Independent confirmation of TUG1-EZH2-MT2A axis lacking
  6. 2020 Medium

    Placed MT2A upstream of the BARD1/BRCA1 module in chemoresistance, expanding its role beyond metal buffering into DNA-repair-associated drug response.

    Evidence Co-IP, immunofluorescence co-localization, and BARD1 rescue of Oxaliplatin resistance in MT2A-knockdown colorectal cancer cells

    PMID:32638210

    Open questions at the time
    • Mechanism by which MT2A stabilizes BARD1/BRCA1 not defined
    • Whether metal binding is required for this interaction untested
  7. 2021 Medium

    Identified MT2A as a negative regulator of NF-κB driving apoptosis and G2/M arrest in AML cells, broadening its tumor-suppressive signaling repertoire.

    Evidence Lentiviral overexpression and shRNA knockdown with flow cytometry and Western blotting of NF-κB components in HL60 cells

    PMID:34220318

    Open questions at the time
    • Direct molecular link between MT2A and IκB-α/NF-κB not established
    • Single cell line
  8. 2022 Medium

    Connected MT2A to Hippo and erythroid-differentiation pathways, showing it activates the MST1/LATS2/YAP1 axis to suppress tumor growth and acts downstream of CD69 via p38MAPK regulation.

    Evidence Overexpression/knockdown with Hippo phosphorylation Westerns and MST1/2 inhibitor rescue plus xenografts (Hippo); MT2A knockdown with p38MAPK inhibition and erythroid readouts (CD69)

    PMID:35642057 PMID:35643179

    Open questions at the time
    • How MT2A activates upstream Hippo kinases is unknown
    • Whether metal buffering mediates these signaling effects untested
  9. 2023 Medium

    Resolved transcriptional control of MT2A, establishing an HSF1 activating input opposed by RING1-mediated HSF1 degradation, and a parallel GHET1-EZH2/LSD1 chromatin-silencing route.

    Evidence Co-IP, ChIP for HSF1 on MT2A promoter, proteasome inhibition and MT2A rescue (RING1-HSF1); RIP, pull-down, histone-mark ChIP, knockdown rescue (GHET1)

    PMID:37548351 PMID:37797799

    Open questions at the time
    • Signals that toggle these activating versus silencing inputs unknown
    • Single-lab studies in distinct cell contexts
  10. 2025 High

    Defined MT2A's role in copper handling and cuproptosis, identifying a druggable site (Lys-31), copper-dependent mitochondrial translocation, and a tetrameric-PKM2 interaction linking metal buffering to metabolic and protective outcomes.

    Evidence HuProt microarray and SPR with site mutagenesis and Parkinson's model (artesunate/Lys-31); mitochondrial proteomics and Co-IP with PKM2; copper-overload and ischemia gain-of-function models

    PMID:39915841 PMID:40754045 PMID:41211480

    Open questions at the time
    • How copper ions trigger mitochondrial translocation mechanistically unresolved
    • Relationship between zinc and copper buffering functions of the same protein unclear

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved how a single metal-buffering protein integrates its many partner-specific and pathway-specific outputs into a unified rule governing cell fate across tissues.
  • No structural model of MT2A in complex with any partner
  • No unified accounting of when MT2A is pro- versus anti-apoptotic
  • Quantitative contribution of metal buffering versus direct protein interactions undefined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Localization
GO:0005739 mitochondrion 2 GO:0005764 lysosome 1
Pathway
R-HSA-1430728 Metabolism 3 R-HSA-162582 Signal Transduction 3 R-HSA-5357801 Programmed Cell Death 3

Evidence

Reading pass · 16 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2015 HMBOX1 physically interacts with MT2A (identified by yeast two-hybrid and confirmed by co-immunoprecipitation), and this interaction elevates intracellular free zinc levels; knockdown of MT2A abolishes HMBOX1-mediated zinc elevation, anti-apoptotic, and pro-autophagic effects in vascular endothelial cells. Yeast two-hybrid, co-immunoprecipitation, MT2A knockdown with phenotypic readout, zinc chelation rescue experiment Scientific reports Medium 26456220
2017 XAF1 directly interacts with MT2A and facilitates its lysosomal degradation, leading to elevation of free intracellular zinc, subsequent p53 activation, and XIAP inactivation, thereby switching stress response toward apoptosis. A MT2A-binding-deficient XAF1 mutant fails to elevate free zinc, confirming the interaction is required for this mechanism. Conversely, MT2A destabilizes XAF1 via the lysosomal pathway under cytostatic stress, establishing mutual antagonism. Co-immunoprecipitation, domain-mutant XAF1 (binding-deficient), lysosomal degradation assays, intracellular zinc measurement, xenograft models, loss-of-function experiments Proceedings of the National Academy of Sciences of the United States of America High 28507149
2006 Recombinant human MT2A expressed in E. coli retains intact metal-binding ability, hydroxyl radical scavenging ability, and protective activity against DNA damage caused by UVC radiation, establishing these as intrinsic biochemical activities of the protein. In vitro biochemical assays: metal-binding, hydroxyl radical scavenging, DNA damage protection assay with UVC-irradiated purified recombinant protein Protein expression and purification Medium 17224279
2014 EOLA1 interacts with MT2A (identified by yeast two-hybrid screening), and MT2A plays a key role in LPS-induced IL-6 expression in HUVECs; EOLA1 acts as a negative regulator of LPS response by regulating MT2A activity. Yeast two-hybrid, knockdown of EOLA1 and MT2A with IL-6 and apoptosis readouts in HUVECs Molecular and cellular biochemistry Medium 24916366
2015 EOLA1 association with MT2A negatively regulates LPS-induced VCAM-1 expression in ECV304 endothelial cells; MT2A knockdown reduces LPS-induced VCAM-1 production, placing MT2A as a positive regulator of VCAM-1 in this inflammatory pathway. Knockdown of EOLA1 and MT2A, overexpression of EOLA1, VCAM-1 measurement in LPS-stimulated ECV304 cells International journal of inflammation Low 26881174
2020 MT2A interacts with BARD1 and BRCA1 (confirmed by co-immunoprecipitation and co-localization by immunofluorescence) and positively regulates BARD1/BRCA1 levels; BARD1 overexpression partially restores Oxaliplatin resistance lost upon MT2A knockdown in colorectal cancer cells, placing MT2A upstream of BARD1/BRCA1 in chemoresistance. Co-immunoprecipitation, immunofluorescence co-localization, MT2A knockdown and overexpression with viability/resistance readouts, BARD1 rescue experiment Cell biochemistry and biophysics Medium 32638210
2022 MT2A overexpression promotes phosphorylation of MST1, LATS2, and YAP1, thereby activating the Hippo signaling pathway and inhibiting colorectal cancer cell proliferation and liver metastasis; specific MST1/2 inhibitors abolish this phosphorylation and rescue proliferation, placing MT2A upstream of the MST1/LATS2/YAP1 axis. Western blotting for Hippo pathway phosphorylation, MT2A overexpression and knockdown in cells, xenograft and liver metastasis animal models, MST1/2 inhibitor rescue experiment, immunohistochemistry Cancer cell international Medium 35642057
2021 MT2A overexpression in HL60 AML cells induces apoptosis and G2/M cell cycle arrest associated with downregulation of p-IκB-α and cyclin D1 and upregulation of IκB-α, placing MT2A as a negative regulator of the NF-κB pathway; MT2A knockdown conversely increases proliferation. Lentiviral MT2A overexpression, shRNA knockdown, flow cytometry, Western blotting for NF-κB pathway components, cell viability assays in HL60 cells International journal of medical sciences Medium 34220318
2022 CD69 inhibits activin A-induced erythroid differentiation via MT2A; activin A reduces MT2A expression through p38MAPK, and MT2A knockdown reduces CD69's ability to suppress erythroid marker expression and sensitization to imatinib in K562 CML cells, placing MT2A downstream of CD69 in this differentiation pathway. MT2A knockdown, CD69 overexpression, p38MAPK inhibition (SB203580), erythroid marker expression, growth inhibition and apoptosis assays in K562 and BCR-ABL+ CD34+ cells Experimental cell research Medium 35643179
2023 HSF1 binds the MT2A promoter to transcriptionally activate MT2A; RING1 (an E3 ubiquitin ligase) binds HSF1 and induces its proteasome-dependent degradation, thereby suppressing MT2A transcription and causing cell cycle arrest and apoptosis in breast cancer cells. Ectopic MT2A expression rescues the anti-proliferative effects of RING1, placing MT2A downstream of the RING1-HSF1 axis. Co-immunoprecipitation (RING1-HSF1), ChIP (HSF1 on MT2A promoter), proteasome inhibitor experiments, MT2A ectopic overexpression rescue, xenograft models, Western blotting Experimental cell research Medium 37797799
2025 Artesunate (AS) directly binds MT2A (validated by HuProt 20K human proteome microarray and surface plasmon resonance), upregulates MT2A in astrocytes, reduces intracellular Cu2+ levels, and inhibits cuproptosis (FDX1, CTR1, lip-DLAT regulation); MT2A-Lys-31 is identified as a key functional site. MT2A knockdown abolishes AS-mediated neuroprotection and copper reduction. Human proteome microarray (HuProt 20K), surface plasmon resonance (SPR), MT2A knockdown, site-directed mutagenesis (Lys-31), copper ion measurements, cuproptosis marker assessment, in vivo 6-OHDA Parkinson's disease model Pharmacological research High 40754045
2025 Under hypoxia, MT2A translocates to mitochondria in a copper-ion-dependent manner and interacts with the tetrameric form of PKM2 to maintain PKM2 enzymatic activity, thereby promoting glycolysis and oxidative phosphorylation in breast cancer cells. Mitochondrial proteomics, co-immunoprecipitation (MT2A-PKM2 tetrameric form), subcellular fractionation, copper ion manipulation, glycolysis and OXPHOS functional assays, MT2A knockdown with tumor growth readout Cell insight Medium 41211480
2025 MT2A overexpression in HUVECs alleviates copper overload (CPO)-induced mitochondrial dysfunction and cytotoxicity; in a rat chronic ischemia model, MT2A overexpression reduces DLAT accumulation and mitochondrial abnormalities and promotes angiogenesis, establishing a copper-mitochondria regulatory mechanism for MT2A in ischemic angiogenesis. MT2A overexpression in HUVECs (in vitro copper overload model), rat 2VO+EMS in vivo model, mitochondrial morphology and activity assays, CD31 immunostaining, cuproptosis marker analysis (DLAT, FDX1, SDHB), cerebral blood perfusion measurement Journal of translational medicine Medium 39915841
2024 MT2A overexpression in hypoxia-exposed dorsal root ganglion neurons inhibits apoptosis by inactivating p38 MAPK, and MT2A co-localizes with neurons (but not microglia or astrocytes) in rat DRG tissues, establishing MT2A as a neuronal anti-apoptotic regulator acting through the p38 MAPK pathway in neurogenic intermittent claudication. Lentivirus-mediated MT2A overexpression in primary neurons, hypoxia-induced cell damage model, co-localization immunostaining, apoptosis assays, p38 MAPK activation measurements, rat cauda equina compression model Human cell Medium 38546949
2019 TUG1 lncRNA recruits EZH2 to the MT2A promoter to epigenetically suppress MT2A transcription; ChIP experiments confirmed EZH2 binding to the MT2A promoter, and TUG1 knockdown reduced this binding, de-repressing MT2A expression. [NOTE: This paper was subsequently retracted (PMID:34992381), so this finding should be treated with caution.] RNA immunoprecipitation (RIP), chromatin immunoprecipitation (ChIP), TUG1 knockdown, MT2A expression analysis OncoTargets and therapy Low 30787623
2023 GHET1 lncRNA recruits EZH2 and LSD1 to the MT2A promoter, reducing H3K27me3 and H3K4me2 occupancy to epigenetically suppress MT2A transcription; MT2A knockdown reverses GHET1 inhibition of trophoblast phenotypes, establishing MT2A as a downstream effector of GHET1 in extravillous trophoblast biology. RNA immunoprecipitation (RIP), RNA pull-down, chromatin immunoprecipitation (ChIP) for histone marks, MT2A knockdown rescue experiments, proliferation and migration assays Molecular reproduction and development Medium 37548351

Source papers

Stage 0 corpus · 40 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2005 Novel -209A/G MT2A polymorphism in old patients with type 2 diabetes and atherosclerosis: relationship with inflammation (IL-6) and zinc. Biogerontology 71 16518702
2015 HMBOX1 interacts with MT2A to regulate autophagy and apoptosis in vascular endothelial cells. Scientific reports 59 26456220
2007 The +838 C/G MT2A polymorphism, metals, and the inflammatory/immune response in carotid artery stenosis in elderly people. Molecular medicine (Cambridge, Mass.) 46 17622311
2018 Arsenic metabolites; selenium; and AS3MT, MTHFR, AQP4, AQP9, SELENOP, INMT, and MT2A polymorphisms in Croatian-Slovenian population from PHIME-CROME study. Environmental research 34 30612060
2019 Overexpressed long noncoding RNA TUG1 affects the cell cycle, proliferation, and apoptosis of pancreatic cancer partly through suppressing RND3 and MT2A. OncoTargets and therapy 33 30787623
2017 Identification of XAF1-MT2A mutual antagonism as a molecular switch in cell-fate decisions under stressful conditions. Proceedings of the National Academy of Sciences of the United States of America 30 28507149
2022 Metallothionein 2A (MT2A) controls cell proliferation and liver metastasis by controlling the MST1/LATS2/YAP1 signaling pathway in colorectal cancer. Cancer cell international 29 35642057
2011 Distributions of interleukin-6 (IL-6) promoter and metallothionein 2A (MT2A) core promoter region gene polymorphisms and their associations with aging in Turkish population. Archives of gerontology and geriatrics 24 21277639
2023 Single-cell transcriptome analysis revealing the intratumoral heterogeneity of ccRCC and validation of MT2A in pathogenesis. Functional & integrative genomics 23 37713131
2016 Metallothionein MT2A A-5G Polymorphism as a Risk Factor for Chronic Kidney Disease and Diabetes: Cross-Sectional and Cohort Studies. Toxicological sciences : an official journal of the Society of Toxicology 22 27122239
2019 MT2A is an early predictive biomarker of response to chemotherapy and a potential therapeutic target in osteosarcoma. Scientific reports 21 31444479
2014 EOLA1 protects lipopolysaccharide induced IL-6 production and apoptosis by regulation of MT2A in human umbilical vein endothelial cells. Molecular and cellular biochemistry 20 24916366
2006 High-yield expression in Escherichia coli of soluble human MT2A with native functions. Protein expression and purification 18 17224279
2020 Polymorphisms in MMP-1, MMP-2, MMP-7, MMP-13 and MT2A do not contribute to breast, lung and colon cancer risk in polish population. Hereditary cancer in clinical practice 14 32765800
2025 MT2A promotes angiogenesis in chronically ischemic brains through a copper-mitochondria regulatory mechanism. Journal of translational medicine 11 39915841
2020 MT2A Promotes Oxaliplatin Resistance in Colorectal Cancer Cells. Cell biochemistry and biophysics 11 32638210
2022 Bioreduction and bioremoval of hexavalent chromium by genetically engineered strains (Escherichia coli MT2A and Escherichia coli MT3). World journal of microbiology & biotechnology 10 35075546
2021 Effect of MT2A on apoptosis and proliferation in HL60 cells. International journal of medical sciences 10 34220318
1987 A 5' flanking region of the metallothionein, MT2A, gene identifies two moderately frequent RFLPs. Nucleic acids research 9 2881275
2015 EOLA1 Inhibits Lipopolysaccharide-Induced Vascular Cell Adhesion Molecule-1 Expression by Association with MT2A in ECV304 Cells. International journal of inflammation 6 26881174
2025 Artesunate alleviates Parkinson's disease by targeting astrocyte MT2A to attenuate dopamine neuronal cuproptosis. Pharmacological research 5 40754045
2022 Activin A downregulates the CD69-MT2A axis via p38MAPK to induce erythroid differentiation that sensitizes BCR-ABL-positive cells to imatinib. Experimental cell research 5 35643179
2022 Blood RNA Sequencing Indicates Upregulated BATF2 and LY6E and Downregulated ISG15 and MT2A Expression in Children with Autism Spectrum Disorder. International journal of molecular sciences 4 36077244
2024 Compensatory upregulation of MT2A alleviates neurogenic intermittent claudication through inhibiting activated p38 MAPK-mediated neuronal apoptosis. Human cell 3 38546949
2023 RING induces cell cycle arrest and apoptosis in human breast cancer cells by regulating the HSF1/MT2A axis. Experimental cell research 3 37797799
2022 ptk2 and mt2a Genes Expression in Gastritis and Gastric Cancer Patients with Helicobacter pylori Infection. Canadian journal of gastroenterology & hepatology 3 36060520
2017 Time-dependent Expression of MT1A mRNA and MT2A mRNA in the Contused Skeletal Muscle of Rats. Fa yi xue za zhi 3 29231000
2010 [Tea polyphenol inhibits colorectal cancer with microsatellite instability by regulating the expressions of HES1, JAG1, MT2A and MAFA]. Zhong xi yi jie he xue bao = Journal of Chinese integrative medicine 3 20836978
2023 lncRNA GHET1 regulates extravillous trophoblastic phenotype via EZH2/LSD1-mediated MT2A epigenetic suppression in pre-eclampsia. Molecular reproduction and development 2 37548351
2014 Interaction study of arsenic (III and V) ions with metallothionein gene (MT2A) fragment. International journal of biological macromolecules 2 25218889
2025 Hypoxia-induced MT2A-tetrameric PKM2 interaction maintains PKM2 activity in a copper-ion-dependent manner. Cell insight 1 41211480
2023 An Assessment of MT1A (rs11076161), MT2A (rs28366003) and MT1L (rs10636) Gene Polymorphisms and MT2 Concentration in Women with Endometrial Pathologies. Genes 1 36981043
2026 MT2A buffering exhaustion marks cuproptosis in severe COVID-19: Multi-omics integration, computational modeling, and experimental validation. Free radical biology & medicine 0 41791684
2026 MT2A-Mediated Regulation of Cuproptosis in Human Dental Pulp Cells Modulates Macrophage M1 Polarization in Pulpitis. International dental journal 0 41863975
2023 Human Infrapatellar Fat Pad Mesenchymal Stem Cell-Derived Extracellular Vesicles Inhibit Fibroblast Proliferation by Regulating MT2A to Reduce Knee Arthrofibrosis. Stem cells international 0 37091533
2021 Overexpressed Long Noncoding RNA TUG1 Affects the Cell Cycle, Proliferation, and Apoptosis of Pancreatic Cancer Partly Through Suppressing RND3 and MT2A [Retraction]. OncoTargets and therapy 0 34992381
2020 Corrigendum to "EOLA1 Inhibits Lipopolysaccharide-Induced Vascular Cell Adhesion Molecule-1 Expression by Association with MT2A in ECV304 Cells". International journal of inflammation 0 33062250
2011 [Immunogenic properties and molecular-genetic features of native and inactivated Chlamydia trachomatis MT-2A serovariant D strain]. Zhurnal mikrobiologii, epidemiologii i immunobiologii 0 22145350
2008 [Biological and immunogenic characteristics of Chlamydia trachomatis strain MT-2A (serovar D) and its use for development of experimental inactivated vaccine]. Zhurnal mikrobiologii, epidemiologii i immunobiologii 0 18595462
2006 [Expression and purification of human MT-2a fusion protein in prokaryotic cells and preparation of its antiserum]. Xi bao yu fen zi mian yi xue za zhi = Chinese journal of cellular and molecular immunology 0 16806025

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