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Showing MTARC1MOSC1 is a alias.

MTARC1

Mitochondrial amidoxime-reducing component 1 · UniProt Q5VT66

Length
337 aa
Mass
37.5 kDa
Annotated
2026-06-10
21 papers in source corpus 16 papers cited in narrative 16 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 6/6 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

MTARC1 (mARC1) is a molybdenum cofactor-containing N-reductive enzyme of the outer mitochondrial membrane that has emerged as a hepatic regulator of lipid droplet homeostasis and a genetically validated protective target in fatty liver disease (PMID:23086957, PMID:37122688, PMID:41641916). It is a signal-anchored protein with an N(in)-C(out) orientation, exposing its C-terminal catalytic domain to the cytosol and assembling into high-oligomeric complexes (PMID:23086957). Its catalytic activity derives from a molybdenum cofactor coordinated by Cys-273; mARC1 reduces N-hydroxylated and N-oxygenated substrates such as benzamidoxime and N-hydroxyurea, and converts nitrite to nitric oxide, operating in an electron-transfer chain with cytochrome b5 and NADH cytochrome b5 reductase (PMID:24500710, PMID:24423752, PMID:30397129, PMID:39397364). In hepatocytes, mARC1 loss reduces triglyceride and neutral lipid accumulation while enhancing fatty acid β-oxidation and lipid secretion, lowering mitochondrial superoxide and ferroptosis under lipotoxic stress and attenuating steatosis, inflammation, and fibrosis across MASH/MASLD models (PMID:37122688, PMID:39716370, PMID:39927988, PMID:41527487). Mechanistically, MTARC1 deficiency post-transcriptionally upregulates the glycerophospholipid biosynthetic enzymes CEPT1 and PEMT, remodeling lipid droplet phospholipid composition to reduce droplet size and promote degradation via lipolysis and lipophagy, an effect reversed by CEPT1 or PEMT knockdown (PMID:41641916). The clinically protective p.A165T variant acts primarily by destabilizing the protein and reducing its levels without altering the fold or active-site architecture, and rare loss-of-function variants are similarly hepatoprotective (PMID:35560545, PMID:38340654, PMID:38437227). mARC1 loss also suppresses proliferation and migration in hepatocellular carcinoma models (PMID:41644117).

Mechanistic history

Synthesis pass · year-by-year structured walk · 9 steps
  1. 2012 High

    Established where mARC1 resides and how it is targeted, defining it as an outer mitochondrial membrane protein with a cytosol-facing catalytic domain.

    Evidence Subcellular fractionation, protease protection, truncation constructs, and ATP/membrane-potential-dependent import assays

    PMID:23086957

    Open questions at the time
    • Functional consequence of high-oligomeric assembly not defined
    • No catalytic substrate identified at this stage
  2. 2014 High

    Defined the catalytic chemistry of mARC1 as molybdenum-dependent N-reduction and nitrite-to-NO conversion within a cytochrome b5/b5 reductase electron-transfer chain.

    Evidence In vitro reconstitution with recombinant protein, C273A active-site mutagenesis, tungsten cofactor substitution, and benzamidoxime kinetics

    PMID:24423752 PMID:24500710

    Open questions at the time
    • Physiological substrates in vivo not established
    • A165T showed no kinetic alteration, leaving its disease mechanism unexplained
  3. 2018 High

    Provided the structural basis of catalysis, placing mARC1 in the MOSC domain superfamily and defining molybdenum cofactor coordination and paralog-distinguishing active-site residues.

    Evidence High-resolution X-ray crystallography with active-site interpretation

    PMID:30397129

    Open questions at the time
    • No structure of substrate-bound complex
    • Mechanism of partner electron-transfer coupling not resolved structurally
  4. 2023 High

    Connected mARC1 to hepatic lipid metabolism by showing hepatocyte-specific knockdown lowers liver triglycerides while raising secretion and altering ketogenesis.

    Evidence GalNAc-siRNA knockdown in a NASH mouse model and primary human hepatocytes with metabolic readouts and RNA-seq

    PMID:37122688

    Open questions at the time
    • Molecular link between N-reductase activity and lipid handling unresolved
    • Increased plasma triglyceride consequence not mechanistically explained
  5. 2024 Medium

    Resolved how the protective p.A165T variant works—by destabilizing the protein and causing aberrant localization rather than altering catalysis—and showed loss-of-function is hepatoprotective in humans.

    Evidence Variant crystallography, protein stability reporter assays across cell lines, knock-in/KO mouse models, localization studies, and exome-wide association (n=540,000)

    PMID:35560545 PMID:38340654 PMID:38437227

    Open questions at the time
    • Mouse-human divergence (Marc2 dominance) complicates modeling
    • Degradation pathway clearing the unstable variant not identified
  6. 2024 High

    Demonstrated that mARC1 loss reduces hepatic lipid burden by enhancing β-oxidation and lowers oxidative stress, ferroptosis, and fibrosis across multiple disease models.

    Evidence siRNA knockdown and global/conditional KO in MASH/MASLD mouse models and primary human hepatocytes with radiolabeled β-oxidation, bioenergetics, mitochondrial superoxide assays, multi-omics, and UK Biobank metabolomics

    PMID:38619429 PMID:38696369 PMID:39716370 PMID:39927988

    Open questions at the time
    • Allele-specific effect (risk-allele homozygotes) not fully mechanistically dissected
    • Direct enzymatic substrate driving lipid phenotype not identified
  7. 2024 Medium

    Refined substrate specificity by establishing mARC1 as the dominant reductase inactivating N-hydroxyurea, distinguishing it from mARC2.

    Evidence In vitro reductive assays with recombinant mARC1 and mARC2 plus in vivo metabolic studies

    PMID:39397364

    Open questions at the time
    • Single lab
    • Broader endogenous N-hydroxylated substrate repertoire not mapped
  8. 2026 High

    Defined the mechanistic axis linking mARC1 to lipid droplet turnover: MTARC1 deficiency post-transcriptionally upregulates CEPT1/PEMT, remodeling droplet phospholipids to drive lipolysis and lipophagy.

    Evidence Global/liver-specific KO mice with epistatic Cept1/Pemt/Pnpla2/Lipa inhibition, multi-omics, and LD size quantification with rescue/reversal experiments

    PMID:41527487 PMID:41641916

    Open questions at the time
    • Mechanism by which mARC1 controls CEPT1/PEMT post-transcriptionally is unknown
    • Whether this axis requires mARC1 catalytic activity is not established
  9. 2026 Medium

    Extended mARC1 function to oncology, showing its loss suppresses HCC proliferation, migration, and lipid accumulation in vitro and in xenografts.

    Evidence siRNA and CRISPR-Cas9 KO across HCC cell lines, β-oxidation and migration assays, xenograft model, and proteomics

    PMID:41644117

    Open questions at the time
    • Single lab
    • Direct molecular driver of the anti-proliferative effect not defined

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unknown how mARC1's molybdenum-dependent N-reductase catalysis mechanistically connects to its control of CEPT1/PEMT and lipid droplet homeostasis, and whether the metabolic phenotype requires catalytic turnover or merely protein presence.
  • No identified endogenous substrate links catalysis to lipid phenotype
  • Post-transcriptional control of CEPT1/PEMT is undefined
  • Causal role of catalysis versus scaffolding in lipid handling unresolved

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0016491 oxidoreductase activity 4 GO:0140098 catalytic activity, acting on RNA 3
Localization
GO:0005739 mitochondrion 2
Pathway
R-HSA-1430728 Metabolism 3

Evidence

Reading pass · 16 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2012 mARC1 is a signal-anchored protein of the outer mitochondrial membrane with an N(in)-C(out) membrane orientation; the N-terminal transmembrane helix is sufficient for mitochondrial targeting, the N-terminal targeting signal acts as a supportive receptor, membrane integration is membrane-potential-independent but requires external ATP, and the protein assembles into high-oligomeric complexes. The C-terminal catalytic domain is exposed to the cytosol. Subcellular fractionation, protease protection assays, deletion/truncation constructs, live-cell imaging, mitochondrial import assays with ATP depletion and membrane-potential uncouplers The Journal of Biological Chemistry High 23086957
2014 Human mARC1 catalyzes reduction of nitrite to nitric oxide (NO) through its molybdenum cofactor; this activity requires Cys-273 (molybdenum-coordinating residue) as the C273A mutation abolishes NO formation. Replacement of molybdenum with tungsten also abolishes NO formation. mARC1 generates NO from nitrite in an electron transfer chain with NADH, cytochrome b5, and NADH-dependent cytochrome b5 reductase, with rate increasing ~3-fold at pH 6.5 vs 7.5. In vitro nitrite reduction assay with recombinant protein; active-site mutagenesis (C273A); tungsten substitution of molybdenum; lentiviral mARC1 expression in HEK cells with NO detection; Km/Vmax determination The Journal of Biological Chemistry High 24500710
2014 mARC1 functions as part of an N-reductive enzyme system together with cytochrome b5 type B and NADH cytochrome b5 reductase to reduce N-hydroxylated compounds (benzamidoxime). SNP variants in MARC1 encoding A165T showed no altered kinetic parameters in benzamidoxime N-reduction, while multiple simultaneous amino acid substitutions reduced N-reductive activity ~5-fold. Recombinant protein expression in E. coli; in vitro steady-state enzyme kinetics with benzamidoxime; molybdenum quantification by ICP-MS; pyrosequencing-based genotyping Drug Metabolism and Disposition Medium 24423752
2018 Crystal structure of human mARC1 was solved at high resolution, revealing the coordination geometry of the molybdenum cofactor (Moco), identifying two key active-site residues that distinguish mARC paralogs, and demonstrating that mARC1 belongs to the MOSC domain superfamily. The structure defines the catalytic mechanism for reduction of N-oxygenated compounds and provides evidence for an evolutionary relationship to the xanthine oxidase superfamily. X-ray crystallography (high-resolution crystal structure); structural comparison with in silico domain predictions; functional interpretation of active-site architecture Proceedings of the National Academy of Sciences of the United States of America High 30397129
2022 Crystal structure of the mARC1 p.A165T variant protein at near-atomic resolution shows that this clinically protective variant does not alter the overall protein fold or active-site architecture compared to wildtype mARC1. X-ray crystallography of variant protein; structural comparison to wildtype crystal structure Hepatology Communications Medium 35560545
2023 Hepatocyte-specific mARC1 knockdown (via GalNAc-siRNA in a NASH mouse model) reduced hepatic triglyceride accumulation but increased plasma triglycerides; in primary human hepatocytes, mARC1 knockdown decreased lipid accumulation and increased triglyceride secretion. mARC1 knockdown also decreased secretion of 3-hydroxybutyrate (a β-oxidation marker) in vitro and in vivo, implicating mARC1 in hepatic lipid metabolism and ketogenesis. GalNAc-siRNA hepatocyte-specific knockdown in GAN-diet NASH mouse model; primary human hepatocyte in vitro model with siRNA KD; metabolic readouts (triglycerides, 3-hydroxybutyrate); RNA-sequencing pathway analysis JHEP Reports High 37122688
2024 The protective MTARC1 p.A165T variant causes dramatically reduced protein stability of mARC1 (assessed by protein stability reporter system in multiple cell lines and in mouse liver), without altering mRNA levels. Multiple substitutions at position A165 (A165S, A165N, A165V, A165G, A165D) similarly reduced stability, indicating A165 is essential for mARC1 protein stability. Protein stability reporter system in multiple cell lines; murine knock-in model (A168T equivalent); Western blot; mutagenesis of A165 to multiple residues Biochemical and Biophysical Research Communications Medium 38340654
2024 The protective p.A165T substitution causes protein instability and aberrant localization of mARC1 in hepatic cells. Novel rare putative loss-of-function MARC1 variants identified by exome-wide association study show a phenotype similar to p.A165T/p.M187K variants, consistent with loss-of-function being hepatoprotective. Marc1 knockout mice, unlike human carriers, do not show protection against hepatic triglyceride accumulation, revealing a divergent physiological role between human and mouse (attributed to Marc2 being the dominant paralog in mouse liver). Exome-wide association study (n=540,000); in vitro expression of recombinant human MARC1 A165T with localization studies in hepatic cells; Marc1 knockout mouse generation; liver phenotype assessment in KO mice on steatogenic diet PLoS Genetics Medium 38437227
2024 Hepatocyte-specific Mtarc1 siRNA knockdown in ob/ob and diet-induced MASH mouse models reduced serum liver enzymes, LDL-cholesterol, liver triglycerides, liver weight, and attenuated liver pathological changes. Multi-omics (metabolomics, proteomics, lipidomics) analysis showed that Mtarc1 knockdown partially restored diet-altered metabolites and lipids. GalNAc-conjugated siRNA hepatocyte-specific knockdown in ob/ob and diet-induced MASH mouse models; multi-omics (metabolomics, proteomics, lipidomics); histology; serum biochemistry Hepatology Communications High 38696369
2024 mARC1 siRNA knockdown in primary human hepatocytes reduced neutral lipid content specifically in cells homozygous for the risk allele (p.A165), and this reduction was mediated by increased fatty acid β-oxidation (measured by radiolabeled tracer). mARC1 knockdown also reduced ferroptosis and reactive oxygen species levels. In human UK Biobank participants, carriers of the rs2642438 minor allele had higher circulating 3-hydroxybutyrate levels, consistent with increased β-oxidation. siRNA knockdown in primary human hepatocytes; radiolabeled fatty acid oxidation assay; Oil-Red O staining; RNA-sequencing and LC-MS proteomics; UK Biobank metabolomics Clinical and Molecular Hepatology High 39716370
2024 mARC1 is the main contributor to reductive biotransformation of N-hydroxyurea (NHU) to urea; in vitro and in vivo evidence establishes that this N-reductive activity is specifically mediated by mARC1 (not mARC2), suggesting mARC1-mediated inactivation as a pharmacological mechanism requiring high doses of hydroxyurea in therapy. In vitro N-reductive assay with recombinant mARC1 and mARC2; in vivo metabolic studies; substrate specificity determination Journal of Medicinal Chemistry Medium 39397364
2024 mARC1 depletion improved cellular bioenergetics and decreased mitochondrial superoxide production in response to lipotoxic stress in cells. The p.A165T variant maintains mitochondrial localization despite lower protein levels. Global or hepatocyte-specific mARC1 deletion in mice reduced liver steatosis and fibrosis in multiple MASH and liver fibrosis models. RNA-seq showed downregulation of extracellular matrix remodeling and collagen formation pathways upon mARC1 loss. mARC1 knockdown/KO in cells with bioenergetics and mitochondrial superoxide assays; global and conditional KO mice on diet-induced MASH models; RNA-seq; plasma lipidomics; histology Hepatology Communications High 39927988
2024 mARC1 modulates lipid accumulation in primary human hepatocytes and primary human adipocytes; mARC1 depletion affects accumulation of distinct lipid species and expression of inflammatory and mitochondrial pathway genes/proteins in both in vitro and in vivo models. Protective MTARC1 variants decrease protein accumulation in overexpression systems (without altering mRNA). A plasma lipid biomarker (Ceramide 22:1) predictive of mARC1 abundance was identified. siRNA knockdown and lentiviral overexpression in primary human hepatocytes, hepatocyte cell lines, and primary human adipocytes; in vivo murine MASH model with GalNAc-siRNA; lipidomics; proteomics; transcriptomics Hepatology Communications Medium 38619429
2026 MTARC1 deficiency post-transcriptionally upregulates glycerophospholipid (GPL) biosynthetic enzymes CEPT1 and PEMT, leading to altered phospholipid composition in lipid droplets (LDs). This phospholipid remodeling reduces LD size, increases surface-to-volume ratio, and thereby enhances LD degradation via lipolysis and lipophagy. Knockdown of CEPT1 or PEMT reversed the hepatoprotective effects of MTARC1 deficiency, establishing an MTARC1-GPL biosynthesis-LD degradation axis. Global and liver-specific Mtarc1 KO mice; genetic inhibition of Pnpla2, Lipa, Pemt, Cept1; multi-omics (biochemical, histological, lipidomics, proteomics); in vitro cell culture mechanistic studies; LD size/number quantification Liver International High 41641916
2026 MTARC1 knockdown in HCC cell lines (Hep3B2, HuH7, HepG2, HepaRG) reduced proliferation; CRISPR-Cas9 KO in Hep3B2 cells decreased neutral lipid accumulation, enhanced β-oxidation, and reduced cell migration. MTARC1 KO xenograft tumors showed reduced volume. Proteomics revealed inhibition of oncogenic pathways and activation of anti-proliferative proteins upon MTARC1 loss. siRNA knockdown in multiple HCC cell lines; CRISPR-Cas9 KO; lipid accumulation assay; β-oxidation assay; migration assay; subcutaneous xenograft mouse model; global proteomics Clinical and Molecular Hepatology Medium 41644117
2026 Mtarc1 KO mice on choline-deficient high-fat diet showed reduced liver steatosis, pro-fibrosis markers, and inflammation. Primary hepatocytes from Mtarc1 KO mice exhibited reduced lipid droplet accumulation, decreased fatty acid uptake, and increased lipid secretion. Metabolomics showed hepatic enrichment of phospholipids in Mtarc1 KO mice. Mtarc1 KO mouse model; CDAHFD-induced MASLD model; primary hepatocyte isolation; lipid droplet imaging; fatty acid uptake assay; lipid secretion assay; untargeted metabolomics Liver International High 41527487

Source papers

Stage 0 corpus · 21 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2014 Nitrite reductase and nitric-oxide synthase activity of the mitochondrial molybdopterin enzymes mARC1 and mARC2. The Journal of biological chemistry 130 24500710
2020 Genome-Wide Association Study for Alcohol-Related Cirrhosis Identifies Risk Loci in MARC1 and HNRNPUL1. Gastroenterology 70 32561361
2012 The mitochondrial amidoxime-reducing component (mARC1) is a novel signal-anchored protein of the outer mitochondrial membrane. The Journal of biological chemistry 63 23086957
2018 Crystal structure of human mARC1 reveals its exceptional position among eukaryotic molybdenum enzymes. Proceedings of the National Academy of Sciences of the United States of America 43 30397129
2023 Hepatocyte mARC1 promotes fatty liver disease. JHEP reports : innovation in hepatology 33 37122688
2024 mARC1 in MASLD: Modulation of lipid accumulation in human hepatocytes and adipocytes. Hepatology communications 15 38619429
2014 Functional characterization of protein variants encoded by nonsynonymous single nucleotide polymorphisms in MARC1 and MARC2 in healthy Caucasians. Drug metabolism and disposition: the biological fate of chemicals 15 24423752
2022 Letter to the editor: The clinically relevant MTARC1 p.Ala165Thr variant impacts neither the fold nor active site architecture of the human mARC1 protein. Hepatology communications 14 35560545
2024 Fatty liver disease protective MTARC1 p.A165T variant reduces the protein stability of MTARC1. Biochemical and biophysical research communications 11 38340654
2024 Downregulation of the MARC1 p.A165 risk allele reduces hepatocyte lipid content by increasing beta-oxidation. Clinical and molecular hepatology 11 39716370
2024 Divergent role of Mitochondrial Amidoxime Reducing Component 1 (MARC1) in human and mouse. PLoS genetics 10 38437227
2024 Liver-specific mitochondrial amidoxime-reducing component 1 (Mtarc1) knockdown protects the liver from diet-induced MASH in multiple mouse models. Hepatology communications 10 38696369
2018 High expression of enhancer RNA MARC1 or its activation by DHT is associated with the malignant behavior in bladder cancer. Experimental cell research 7 29964053
2025 Loss of mitochondrial amidoxime-reducing component 1 (mARC1) prevents disease progression by reducing fibrosis in multiple mouse models of chronic liver disease. Hepatology communications 6 39927988
2025 Polymorphism's MBOAT7 as Risk and MTARC1 as Protection for Liver Fibrosis in MASLD. International journal of molecular sciences 4 40650184
2026 Loss of Mtarc1 Protects Against Steatotic Liver Disease in Mice. Liver international : official journal of the International Association for the Study of the Liver 2 41527487
2024 mARC1 Is the Main Contributor to Metabolic Reduction of N-Hydroxyurea. Journal of medicinal chemistry 1 39397364
2026 A mitochondrial amidoxime-reducing component 1 (mARC1) A168T amino acid substitution does not confer protection from MASH and fibrosis in multiple mouse models of chronic liver disease. The Biochemical journal 0 41428769
2026 MTARC1 Inactivation Remodels Lipid Droplets to Protect Against Metabolic Fatty Liver Disease. Liver international : official journal of the International Association for the Study of the Liver 0 41641916
2026 MTARC1 p.A165 ablation reduces hepatocellular carcinoma aggressiveness in vitro and in vivo. Clinical and molecular hepatology 0 41644117
2024 Generation of an induced pluripotent stem cell (iPSC) line (EXSURGi001-A) from a patient homozygous for the p.Ala165Thr mutation in the MTARC1 gene. Stem cell research 0 39094505

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