Affinage

DBR1

Lariat debranching enzyme · UniProt Q9UK59

Length
544 aa
Mass
61.6 kDa
Annotated
2026-06-09
26 papers in source corpus 18 papers cited in narrative 17 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

DBR1 is the sole RNA lariat debranching enzyme in eukaryotic cells, a binuclear metallophosphoesterase that hydrolyzes the 2'-5' phosphodiester bond at the branchpoint of excised intron lariats and is thereby the rate-limiting step in lariat turnover (PMID:16275784, PMID:10982890, PMID:38816363). Catalysis proceeds by a two-metal mechanism in which a Cys11-Xaa-His13 motif coordinates the active-site metals and His86 serves as a general acid that protonates the O2' leaving group, with structures of substrate- and product-bound enzyme defining how Dbr1 specifically recognizes 2',5'- but not 3',5'-linkages (PMID:25123664, PMID:27765821, PMID:36484984). Although early work scored the enzyme as manganese-dependent, the biological cofactors are Fe2+ in the β-pocket and Zn2+ in the α-pocket, and iron chelation in human cells raises lariat levels, establishing Fe as physiologically relevant (PMID:16275784, PMID:27930312, PMID:35459748, PMID:37507019). In human cells DBR1 is predominantly nuclear, is recruited to the branchpoint via the intron-binding protein AQR, and preferentially acts on substrates bearing canonical U2 motifs; its debranching activity is required for spliceosome recycling, since loss of DBR1 causes ~20-fold lariat accumulation, prolonged retention of spliceosomal components on lariats, and increased exon skipping (PMID:38816363, PMID:28504715). The disordered C-terminal domain of human Dbr1 reduces aggregation and provides a docking site for the noncatalytic partner TTDN1 (MPLKIP), which both stabilizes DBR1 and stimulates its catalytic efficiency ~19-fold (PMID:37507019, PMID:37800682). Beyond splicing, DBR1 supports innate antiviral immunity: when lariats accumulate they reach the cytosol and form Alu-derived dsRNA hairpins, drive proteasomal degradation of the stress-granule scaffolds G3BP1/2, and blunt PKR and other dsRNA sensors, with PKR ablation abolishing the antiviral effect and Dbr1-hypomorphic mice showing susceptibility to viral infection [PMID:39636299, PMID:bio_10.1101_2024.12.07.627371]. Inherited DBR1 deficiency (e.g. the I120T variant) lowers DBR1 protein, causes lariat accumulation, and increases susceptibility of patient and stem-cell-derived hindbrain neurons to viral infection, a phenotype rescued by wild-type DBR1 (PMID:39023559).

Mechanistic history

Synthesis pass · year-by-year structured walk · 12 steps
  1. 2000 High

    Establishing that the human gene encodes a bona fide debranching enzyme converted a yeast activity into a defined human enzyme and showed functional conservation across eukaryotes.

    Evidence in vitro debranching assay with recombinant hDBR1 plus cross-species complementation of S. cerevisiae and S. pombe dbr1 nulls

    PMID:10982890

    Open questions at the time
    • No structural or mechanistic detail of the active site
    • Physiological metal cofactor not defined
  2. 2005 High

    Mutagenesis defined the catalytic residue set required for debranching, mapping the functional active site of the enzyme.

    Evidence alanine-scanning of 28 conserved residues with in vitro and in vivo lariat assays in yeast Dbr1

    PMID:16275784

    Open questions at the time
    • Manganese assigned as cofactor, later revised
    • Catalytic roles of individual residues not yet mechanistically assigned
  3. 2014 High

    Crystal structures with branchpoint-mimicking RNA explained substrate recognition and the strict selectivity for 2',5'- over 3',5'-phosphodiester bonds.

    Evidence X-ray structures of apo-Dbr1 and Dbr1–branched RNA complexes with functional variant validation

    PMID:25123664

    Open questions at the time
    • Catalytic chemistry (acid/base, metal roles) not fully resolved
    • Identity of physiological metals not addressed
  4. 2016 High

    Combined structural, kinetic, and reconstitution work redefined Dbr1 as an Fe/Zn binuclear enzyme using a two-metal mechanism with His86 as general acid, correcting the earlier manganese model.

    Evidence Fe-edge anomalous diffraction, metal reconstitution, alanine-scanning, and phosphodiesterase kinetics in E. histolytica and yeast Dbr1

    PMID:27765821 PMID:27930312

    Open questions at the time
    • Whether human Dbr1 uses the same metals not yet shown in cells
    • In-cell metal occupancy unknown
  5. 2022 High

    Iron chelation in human cells raised cellular lariat levels, establishing Fe as a biologically important cofactor rather than an in vitro artifact, and product-bound structures illuminated the kinetic mechanism.

    Evidence ICP-MS, apoenzyme reconstitution kinetics, deferoxamine treatment of human lymphoblasts; product-bound PS-bRNA crystal structure

    PMID:35459748 PMID:36484984

    Open questions at the time
    • Direct measurement of Fe occupancy of Dbr1 in human cells not shown
    • Extent of product inhibition in vivo unknown
  6. 2023 High

    Identification of TTDN1/MPLKIP as a direct partner that binds the disordered C-terminal domain, stabilizes DBR1, and boosts catalytic efficiency linked Dbr1 regulation to a defined cofactor protein.

    Evidence NMR characterization, pulldown binding, reconstituted kinetics with TTDN1, and interaction proteomics in patient fibroblasts

    PMID:37507019 PMID:37800682

    Open questions at the time
    • Structural basis of TTDN1–Dbr1 interaction not resolved
    • How TTDN1 mechanistically enhances catalysis is unknown
    • MPLKIP stabilization mechanism not reconstituted
  7. 2024 High

    A viable human knockout demonstrated DBR1 is the sole debranching enzyme, defined AQR-mediated recruitment to branchpoints, and established a direct role in spliceosome recycling and suppression of exon skipping.

    Evidence DBR1 KO cell line, co-IP mass spectrometry, ADAR lariat timestamping, transcriptomics, and spliceosome association assays

    PMID:38816363

    Open questions at the time
    • Mechanism by which AQR delivers Dbr1 to the branchpoint not structurally defined
    • Substrate preference determinants only partially mapped
  8. 2024 High

    Knockout, epistasis, and mouse models established a non-splicing role for DBR1 in antiviral immunity, in which lariat accumulation degrades G3BP1/2, impairs stress granules, and blunts PKR.

    Evidence DBR1-deficient cells, PKR knockout epistasis, Dbr1Y17H mice, proteasome inhibition, and antiviral infection assays

    PMID:39636299

    Open questions at the time
    • How lariats trigger G3BP1/2 degradation mechanistically is unresolved
    • Which sensors dominate in different tissues not defined
  9. 2024 High

    Patient-derived cells and neurons established inherited DBR1 deficiency as a cause of lariat accumulation and impaired antiviral immunity, with rescue and gain-of-function lariat transfection proving causality.

    Evidence DBR1 I120T patient fibroblasts, lentiviral WT rescue, hPSC-derived hindbrain neurons, SARS-CoV-2 infection, and exogenous lariat transfection

    PMID:39023559

    Open questions at the time
    • Tissue-specific basis of clinical phenotype not fully explained
    • Quantitative threshold of lariat accumulation needed for disease unknown
  10. 2024 Medium

    Cytosolic accumulation of lariats, including Alu-derived dsRNA hairpins, was shown to chronically desensitize multiple dsRNA sensors, broadening the antiviral mechanism beyond PKR alone.

    Evidence DBR1 KO cells, dsRNA immunofluorescence, antiviral pathway readouts, and HSV-1/influenza/KSHV infections (preprint)

    PMID:bio_10.1101_2024.12.07.627371

    Open questions at the time
    • Preprint, not yet peer-reviewed
    • Relative contribution of MDA5/RIG-I/RNase L vs PKR not quantified
  11. 2017 Medium

    Knockdown studies linked DBR1 to HIV-1 replication, indicating the enzyme is required for forming or resolving a lariat-like structure during reverse transcription.

    Evidence siRNA/shRNA knockdown with cDNA rescue, cell fractionation, 5' RACE, RNase R resistance, and in vitro DBR1 catalytic-mutant control

    PMID:16232320 PMID:24672043 PMID:28931690

    Open questions at the time
    • Direct vs indirect requirement not fully resolved by knockdown
    • Nature of the HIV lariat-like substrate incompletely defined
  12. 2026 Medium

    Interactome and phosphoproteomic profiling placed hDBR1 within RNA quality-control and stress-granule networks and identified regulatory phosphorylation sites.

    Evidence immunopurification mass spectrometry with RNase A treatment and phosphoproteomics

    PMID:41999910

    Open questions at the time
    • Specific interactions not functionally validated
    • Functional consequence of phosphosites unknown

Open questions

Synthesis pass · forward-looking unresolved questions
  • How lariat accumulation is sensed to trigger proteasomal G3BP1/2 degradation, and how Dbr1 activity and recruitment are regulated by phosphorylation and partner proteins in vivo, remain open.
  • No molecular mechanism connecting lariats to G3BP1/2 turnover
  • Functional role of identified phosphosites unestablished
  • Structure of AQR/TTDN1 complexes with Dbr1 unresolved

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140098 catalytic activity, acting on RNA 5 GO:0016787 hydrolase activity 3 GO:0003723 RNA binding 2
Localization
GO:0005829 cytosol 2 GO:0005634 nucleus 1
Pathway
R-HSA-168256 Immune System 3 R-HSA-8953854 Metabolism of RNA 2

Evidence

Reading pass · 17 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2005 Yeast Dbr1 is a manganese-dependent RNA debranching enzyme that cleaves the 2'-5' phosphodiester bonds of lariat introns. Alanine scanning mutagenesis identified 13 conserved residues (His13, Asp40, Arg45, Asp49, Tyr68, Tyr69, Asn85, His86, Glu87, His179, Asp180, His231, His233) required for function in vivo; mutation of Asp40, Asn85, His86, His179, His231, or His233 to alanine abolished or greatly diminished debranching activity in vitro. Dbr1 sediments as a monomer and requires manganese as the metal cofactor. In vitro debranching assays with natural lariat RNAs and synthetic branched RNAs; alanine-scanning mutagenesis of 28 conserved residues; sedimentation analysis Nucleic acids research High 16275784
2000 Human DBR1 (hDBR1) encodes a functional RNA lariat debranching enzyme: recombinant hDBR1 expressed in E. coli showed debranching activity in vitro, and the cDNA complemented both the intron accumulation phenotype of S. cerevisiae dbr1 null mutants and the intron accumulation and slow growth phenotypes of S. pombe dbr1 null mutants. In vitro debranching assay with recombinant protein; interspecies complementation of S. cerevisiae and S. pombe dbr1 null mutants Nucleic acids research High 10982890
2014 Crystal structures of Dbr1 alone and in complex with synthetic RNA compounds mimicking the lariat branchpoint revealed the molecular basis for 2',5'-phosphodiester recognition and explained why Dbr1 lacks activity toward 3',5'-phosphodiester linkages. Functional data on Dbr1 variants confirmed structure-function relationships. X-ray crystallography of apo-Dbr1 and Dbr1–branched RNA complexes; functional assays on Dbr1 variants Nucleic acids research High 25123664
2016 Entamoeba histolytica Dbr1 contains nearly stoichiometric Fe and Zn per polypeptide; Fe partitions primarily to the β-metal pocket and Zn to the α-pocket. Apo-Dbr1 reconstituted with Fe(II)+Zn(II) is active (~3–4 s⁻¹), while Mn(II) or Fe(II) under aerobic conditions yields inactive enzyme. Co-crystal structures of catalytic mutant H91A with 7-mer and 16-mer branched RNAs show a bridging hydroxide positioned for nucleophilic attack of the scissile phosphate. X-ray crystallography (Fe-edge anomalous diffraction); fluorogenic bRNA kinetic assay; metal reconstitution of apoenzyme under aerobic and anaerobic conditions; elemental analysis Proceedings of the National Academy of Sciences of the United States of America High 27930312
2016 Yeast Dbr1 active site analysis established: (i) Cys11 is an essential active-site residue, extending the Cys11-Xaa-His13 motif as the metal-binding element (atypical within the metallophosphoesterase superfamily); (ii) His86 acts as a general acid catalyst that protonates the O2' leaving group of the 2'-5' phosphodiester; (iii) Dbr1 adheres to a two-metal catalytic mechanism. Dbr1 also exhibits vigorous Mn²⁺-dependent phosphodiesterase activity toward bis-p-nitrophenylphosphate, which does not require His86. Alanine-scanning mutagenesis; in vitro phosphodiesterase assay with bis-p-nitrophenylphosphate and branched RNA substrates; structure-activity analysis RNA (New York, N.Y.) High 27765821
2022 Saccharomyces cerevisiae Dbr1 contains stoichiometric Fe and Zn; Fe²⁺ is the most effective cofactor for reconstituting activity in apoenzyme (turnover ~9.2 s⁻¹). Treatment of human lymphoblastoid cells with the iron chelator deferoxamine caused a ~2-fold increase in cellular lariat levels, indicating Fe is an important biological cofactor for Dbr1. Elemental analysis (ICP-MS); apoenzyme reconstitution with Fe²⁺, Zn²⁺, Mn²⁺; fluorogenic bRNA kinetic assay; deferoxamine treatment of human cells with lariat quantification RNA (New York, N.Y.) High 35459748
2022 A product-bound crystal structure of E. histolytica Dbr1 co-crystallized with a phosphorothioate-branched RNA (PS-bRNA) showed in-crystal hydrolysis of the phosphorothioate bond, revealing a product-bound active site state. Dbr1 cleaves phosphorothioate linkages ~10,000-fold more slowly than native phosphate linkages. The structure suggests product inhibition may contribute to the kinetic mechanism. X-ray crystallography of EhDbr1 with PS-bRNA; kinetic comparison of phosphorothioate vs native substrate cleavage Biochemistry High 36484984
2023 Human Dbr1 contains a disordered C-terminal domain (characterized by NMR and sequence analysis) that stabilizes Dbr1 by reducing aggregation but is dispensable for debranching activity. The noncatalytic protein Drn1 and TTDN1 (MPLKIP) directly bind human Dbr1. Addition of TTDN1 to in vitro debranching reactions increases the catalytic efficiency of human Dbr1 19-fold but has no effect on E. histolytica Dbr1, which lacks the disordered C-terminal domain. Dbr1 requires Fe²⁺ for efficient catalysis. The identity of the branchpoint nucleotide affects debranching rates. NMR spectroscopy; in vitro binding assays (pulldown); in vitro debranching kinetics with TTDN1; comparative analysis with E. histolytica Dbr1; sequence analysis The Journal of biological chemistry High 37507019
2023 MPLKIP (TTDN1) interacts with core splicing factors and the lariat debranching protein DBR1 (identified by mass spectrometry-based interaction proteomics). MPLKIP-deficient primary fibroblasts have reduced steady-state DBR1 protein levels, indicating MPLKIP stabilizes DBR1. Mass spectrometry-based interaction proteomics (co-immunoprecipitation); immunoblot of MPLKIP-deficient fibroblasts EMBO molecular medicine Medium 37800682
2024 DBR1 is the sole debranching enzyme in human cells (demonstrated by knockout). The predominantly nuclear Dbr1 preferentially debranches substrates containing canonical U2 binding motifs and shows specificity for particular 5' splice site sequences. Dbr1 is recruited to the branchpoint through the intron-binding protein AQR (identified by co-immunoprecipitation mass spectrometry). DBR1 depletion causes a 20-fold increase in lariats, increases exon skipping, and causes spliceosomal components to remain associated with lariats for longer, demonstrating a role for Dbr1 in spliceosome recycling. DBR1 knockout cell line generation; co-immunoprecipitation mass spectrometry; ADAR-fusion lariat timestamping; transcriptomic analysis; spliceosome association assays Nature communications High 38816363
2024 Accumulation of RNA lariats in human DBR1-deficient cells interferes with stress granule (SG) assembly by promoting proteasome-mediated degradation of G3BP1 and G3BP2. Impaired SG assembly reduces PKR recruitment and activation, impairing antiviral immunity against HSV-1. Genetic ablation of PKR abolished the antiviral effect of DBR1 in vitro. Dbr1Y17H/Y17H mice are susceptible to viral infections and show decreased G3BP1/2 expression and reduced PKR phosphorylation in cells and brain. DBR1-deficient human cell lines; PKR knockout epistasis; Dbr1Y17H/Y17H mouse model; proteasome inhibition; immunoblot for G3BP1/2 and p-PKR; antiviral infection assays The Journal of experimental medicine High 39636299
2024 DBR1 I120T/I120T homozygous fibroblasts from patients with inherited DBR1 deficiency have low DBR1 protein levels and high RNA lariat accumulation. Exogenous WT DBR1 expression in patient-derived DBR1 I120T/I120T fibroblasts and hPSC-derived hindbrain neurons rescued the RNA lariat accumulation phenotype. Expression of exogenous RNA lariats (mimicking DBR1 deficiency) increased susceptibility of WT hindbrain neurons to SARS-CoV-2 infection, indicating that lariat accumulation per se impairs antiviral immunity. Patient fibroblast analysis; lentiviral rescue with WT DBR1; hPSC-derived hindbrain neuron differentiation; SARS-CoV-2 infection assay; exogenous lariat transfection The Journal of experimental medicine High 39023559
2017 DBR1 knockdown in HIV-infected cells inhibits formation of intermediate and full-length HIV-1 cDNA without affecting minus-strand strong-stop cDNA. This inhibition occurs in the nucleus or perinuclear region: when nuclear import of the HIV-1 reverse transcription complex is blocked by a truncated CPSF6, reverse transcription is completed in the cytoplasm in a DBR1-independent manner. In vitro incubation with yeast or human DBR1 (but not catalytically inactive DBR1(N85A)) resolved a lariat-like structure at the 5' end of HIV-1 RNA detected in infected DBR1-knockdown cells. HIV-1 RNA from DBR1-knockdown cells was resistant to RNase R (which degrades linear but not circular/lariat RNAs), supporting formation of a lariat-like structure. shRNA knockdown; cell fractionation; CPSF6 truncation for nuclear import block; 5' RACE; RNase R treatment; in vitro DBR1 treatment with catalytic mutant control Journal of virology Medium 24672043 28931690
2005 siRNA-mediated knockdown of human DBR1 expression (reducing mRNA by ~80%) led to decreases in HIV-1 cDNA and protein production; this effect was reversed by cotransfection of a DBR1 cDNA, confirming specificity of the DBR1 requirement for HIV-1 replication. siRNA knockdown; DBR1 cDNA rescue; HIV-1 cDNA quantification; viral protein detection Retrovirology Medium 16232320
2017 hDBR1 modulates snRNP recycling to affect alternative RNA splicing. Insufficient hDBR1 leads to higher rates of exon skipping confirmed by transcriptomic sequencing. Wild-type p53 and HIF-1 co-regulate hDBR1 expression. Higher hDBR1 expression only affected exon-skipping activity in malignant (not normal) cells. siRNA knockdown; transcriptomic sequencing; metabolite profiling; in vivo tumor models; reporter assays for splicing Oncogene Medium 28504715
2026 hDBR1 physically associates with spliceosome components, intron-turnover factors (including AQR), and RNA quality-control proteins (UPF1, XRN2, DHX29) as identified by immunopurification coupled to mass spectrometry. RNase A treatment identified an RNA-dependent subnetwork enriched for stress-granule proteins and hnRNPs. Phosphoproteomic profiling identified multiple hDBR1 phosphorylation sites, including four residues preferentially detected after RNase treatment. Immunopurification coupled to mass spectrometry (gel-based and on-bead workflows); RNase A treatment; phosphoproteomic profiling Cell stress & chaperones Medium 41999910
2024 In DBR1-null cells, lariats accumulate in the cytosol and dsRNA (formed by inverted Alu element hairpins within introns) becomes enriched. Chronic exposure to accumulated lariats attenuates dsRNA sensors MDA5, RIG-I, RNase L, and PKR, reducing the antiviral response. Lariats are transiently elevated during infection (HSV-1, influenza, KSHV). DBR1 knockout cells; dsRNA immunofluorescence and quantification; antiviral pathway activation assays; infection experiments with HSV-1, influenza, KSHV bioRxivpreprint Medium bio_10.1101_2024.12.07.627371

Source papers

Stage 0 corpus · 26 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2005 Structure-function analysis of yeast RNA debranching enzyme (Dbr1), a manganese-dependent phosphodiesterase. Nucleic acids research 52 16275784
2005 DBR1 siRNA inhibition of HIV-1 replication. Retrovirology 49 16232320
2000 Human RNA lariat debranching enzyme cDNA complements the phenotypes of Saccharomyces cerevisiae dbr1 and Schizosaccharomyces pombe dbr1 mutants. Nucleic acids research 41 10982890
2014 Structural basis of lariat RNA recognition by the intron debranching enzyme Dbr1. Nucleic acids research 40 25123664
2017 Human DBR1 modulates the recycling of snRNPs to affect alternative RNA splicing and contributes to the suppression of cancer development. Oncogene 39 28504715
2014 Impairment of HIV-1 cDNA synthesis by DBR1 knockdown. Journal of virology 28 24672043
2016 Metal dependence and branched RNA cocrystal structures of the RNA lariat debranching enzyme Dbr1. Proceedings of the National Academy of Sciences of the United States of America 24 27930312
2020 Dbr1 functions in mRNA processing, intron turnover and human diseases. Biochimie 22 33038423
2024 SARS-CoV-2 brainstem encephalitis in human inherited DBR1 deficiency. The Journal of experimental medicine 19 39023559
2017 Conformational Changes in the 5' End of the HIV-1 Genome Dependent on the Debranching Enzyme DBR1 during Early Stages of Infection. Journal of virology 15 28931690
2024 The debranching enzyme Dbr1 regulates lariat turnover and intron splicing. Nature communications 14 38816363
2024 Sequestration of DBR1 to stress granules promotes lariat intronic RNAs accumulation for heat-stress tolerance. Nature communications 14 39227617
2016 Mechanistic insights into the manganese-dependent phosphodiesterase activity of yeast Dbr1 with bis-p-nitrophenylphosphate and branched RNA substrates. RNA (New York, N.Y.) 13 27765821
2024 Human DBR1 deficiency impairs stress granule-dependent PKR antiviral immunity. The Journal of experimental medicine 12 39636299
2023 Trichothiodystrophy-associated MPLKIP maintains DBR1 levels for proper lariat debranching and ectodermal differentiation. EMBO molecular medicine 11 37800682
2017 Fluorescent Branched RNAs for High-Throughput Analysis of Dbr1 Enzyme Kinetics and Inhibition. ACS chemical biology 11 28055181
2023 A founder DBR1 variant causes a lethal form of congenital ichthyosis. Human genetics 10 37656279
2022 Metal content and kinetic properties of yeast RNA lariat debranching enzyme Dbr1. RNA (New York, N.Y.) 8 35459748
2023 Activation of human RNA lariat debranching enzyme Dbr1 by binding protein TTDN1 occurs though an intrinsically disordered C-terminal domain. The Journal of biological chemistry 6 37507019
2025 DBR1 orchestrates the fate of lariat RNA: debranching-dependent turnover and function. Nucleic acids research 4 40637223
2022 Crystal Structure of the RNA Lariat Debranching Enzyme Dbr1 with Hydrolyzed Phosphorothioate RNA Product. Biochemistry 3 36484984
1994 Mapping of DBR1 and YPK1 suggests a major revision of the genetic map of the left arm of Saccharomyces cerevisiae Chromosome XI. Genetics 3 7828812
2025 RNA Lariat-Debranching Enzyme (DBR1) Variations in Sabinas Brittle Hair Syndrome Form of Trichothiodystrophy: A Trichothiodystrophy-Causing Gene. The Journal of investigative dermatology 2 40683339
2023 The debranching enzyme Dbr1 regulates lariat turnover and intron splicing. Research square 2 37398028
2026 The human DBR1 interactome reveals coupling between intron lariat turnover, pre-mRNA splicing, and RNA quality control pathways. Cell stress & chaperones 0 41999910
2025 DBR1 Gene Mutation: Pathogenicity in the Homozygous State and Its Phenotype in Two Siblings. Clinical genetics 0 41366235

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