Affinage

SMUG1

Single-strand selective monofunctional uracil DNA glycosylase · UniProt Q53HV7

Length
270 aa
Mass
29.9 kDa
Annotated
2026-06-10
100 papers in source corpus 19 papers cited in narrative 19 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 8/8 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

SMUG1 is a base excision repair DNA glycosylase that initiates antimutational removal of uracil and oxidized pyrimidines, serving as the principal replication-independent backup to UNG for repair of deaminated cytosine (PMID:11483530, PMID:15902269). It excises uracil, 5-hydroxymethyluracil, 5-hydroxyuracil, and 5-formyluracil from DNA, while excluding thymine and analogous cytosine derivatives, and additionally acts as a xanthine-DNA glycosylase (PMID:11526119, PMID:12718543, PMID:18835277). Crystallography and active-site mutagenesis define how SMUG1 reads the lesion: an invasive interaction with double-stranded DNA, a water displacement/replacement strategy that admits C5-substituted uracils but excludes thymine, and defined catalytic and recognition residues (Asn85, His239, Phe98, Asn163, and the Gly87-Met91 loop) (PMID:12820976, PMID:15466595). Pre-steady-state kinetics establish a wedge-based lesion-search mechanism in which catalysis is rate-limited by tight binding to the abasic-site product, a property that lets SMUG1 inhibit AP-site cleavage by APE1 and that distinguishes it from rapid-turnover UNG2 (PMID:17537817, PMID:29051947); under physiological salt SMUG1 is predominantly double-strand specific, explaining its limited capacity to substitute for UNG in single-strand contexts (PMID:22483865). Genetically, SMUG1 and UNG have non-redundant roles in suppressing C-to-T mutagenesis and in preventing genomic accumulation of 5-hydroxymethyluracil, with combined deficiency producing synergistic uracil accumulation and CpG-biased mutational signatures (PMID:15902269, PMID:22447450, PMID:28775312). SMUG1 also processes 5-fluorouracil incorporated into DNA, protecting against its cytotoxicity and supporting replication recovery after drug exposure (PMID:17283124, PMID:23253900), and contributes to immunoglobulin class switching and A:T somatic hypermutation as a backup to UNG (PMID:16407970, PMID:24771041). Beyond DNA, SMUG1 functions in RNA quality control through interaction with the pseudouridine synthase DKC1, governing rRNA maturation and co-transcriptional processing of the telomerase RNA component hTERC required for telomerase biogenesis (PMID:23246433, PMID:31412240). Its 5-hmdU excision is stimulated by UV-DDB, which displaces SMUG1 from abasic products to relieve product inhibition (PMID:36971122).

Mechanistic history

Synthesis pass · year-by-year structured walk · 18 steps
  1. 2001 High

    Established SMUG1 as the major UNG-independent uracil-excision activity in vivo, defining a replication-independent backup arm of cytosine-deamination repair distinct from replication-coupled UNG.

    Evidence Antibody neutralization of glycosylase activity in Ung-/- mouse cells and subcellular fractionation

    PMID:11483530

    Open questions at the time
    • Did not define the full lesion spectrum
    • Did not establish whether the backup role suffices to prevent mutagenesis
  2. 2001 High

    Identified SMUG1 as the enzyme behind HMUDG activity, extending its substrate range beyond uracil to the oxidation product 5-hydroxymethyluracil.

    Evidence Protein purification, mass spectrometry, and recombinant enzyme assays from calf thymus/human SMUG1

    PMID:11526119

    Open questions at the time
    • Did not establish in vivo importance of hmU excision
    • Strandedness preference not resolved
  3. 2003 High

    Resolved the structural basis for substrate discrimination, showing how SMUG1 excludes thymine yet accepts C5-modified uracils via a water displacement mechanism and an unusually invasive dsDNA interaction.

    Evidence X-ray crystallography of SMUG1-DNA base-excision product complex; recombinant rat/human enzyme substrate-specificity assays with antibody neutralization in HeLa extracts

    PMID:12718543 PMID:12820976

    Open questions at the time
    • Catalytic residue assignments not yet functionally tested
    • Kinetics of recognition and product release unresolved
  4. 2004 High

    Defined the catalytic and recognition residues, mechanistically explaining how SMUG1 hydrolyzes the glycosidic bond and reads diverse C5 substituents.

    Evidence Site-directed mutagenesis of hSMUG1 with multi-substrate activity assays and homology modeling

    PMID:15466595

    Open questions at the time
    • Did not address product binding/turnover kinetics
    • Modeling rather than human crystal structure
  5. 2005 High

    Demonstrated that SMUG1 and UNG are genetically non-redundant in suppressing C-to-T transitions and that SMUG1 also defends against oxidative cytosine lesions.

    Evidence Stable siRNA knockdown in mouse fibroblasts, lacZ mutation frequency, Ung-/- double-mutant epistasis, and ionizing-radiation sensitivity

    PMID:15902269

    Open questions at the time
    • Knockdown rather than complete knockout
    • Molecular identity of the relevant oxidative cytosine lesion not defined
  6. 2006 High

    Clarified why SMUG1 contributes little to antibody diversification under physiological conditions despite being able to substitute for UNG when overexpressed.

    Evidence SMUG1 overexpression in DT40 cells and transgenic mice; SHM/class-switch assays and B-cell expression analysis

    PMID:16407970

    Open questions at the time
    • Did not resolve the biochemical basis for the preference for conventional repair
    • Endogenous contribution in Ung-/- not yet quantified
  7. 2007 High

    Showed SMUG1 uniquely processes genomic 5-fluorouracil and that genomic FU incorporation, not uracil excision, drives FU cytotoxicity, linking SMUG1 to chemotherapy response.

    Evidence Gene-targeted UNG/SMUG1/double-deficient cell lines, FU cytotoxicity assays, and genomic FU/uracil measurements

    PMID:17283124

    Open questions at the time
    • Downstream repair steps after FU excision not defined
    • Did not address replication recovery dynamics
  8. 2007 High

    Defined the divergent BER initiation logic of SMUG1 versus UNG2, showing SMUG1 binds AP-site products tightly and inhibits APE1 cleavage while UNG2 promotes rapid turnover.

    Evidence In vitro AP-site binding/cleavage assays, E. coli complementation, and mutagenesis of an AP-site-binding motif

    PMID:17537817

    Open questions at the time
    • Cellular factors relieving product inhibition not identified
    • Physiological consequence of APE1 inhibition unresolved
  9. 2008 High

    Extended SMUG1's catalytic repertoire to xanthine excision and mapped the active-site determinants distinguishing UDG from XDG activity.

    Evidence Mutagenesis of bacterial and human SMUG1, in vitro UDG/XDG assays, and molecular dynamics simulations

    PMID:18835277

    Open questions at the time
    • In vivo relevance of xanthine excision not established
    • Frequency of xanthine lesions in genomic DNA not addressed
  10. 2012 High

    Genetic ablation in mice confirmed SMUG1 as the dominant hmU-excising glycosylase and the major UNG backup for uracil, validating prior biochemical assignments in vivo.

    Evidence Single and double Smug1/Ung knockout mice, tissue extract glycosylase assays, and 5-hmdU cytotoxicity in fibroblasts

    PMID:22447450

    Open questions at the time
    • Did not quantify genomic lesion accumulation
    • Tissue-specific contributions not resolved
  11. 2012 High

    Provided the biochemical explanation for SMUG1's limited role at Ig loci by showing it is predominantly double-strand specific under physiological salt, unlike UNG2 on ssDNA.

    Evidence In vitro UDG assays under varying ionic conditions with ss/dsDNA and AID-hotspot substrates

    PMID:22483865

    Open questions at the time
    • Did not test whether cellular cofactors alter strand preference
    • In vivo strand context at lesion sites not measured
  12. 2012 High

    Revealed an RNA-directed function for SMUG1 in ribosomal RNA quality control via interaction with DKC1, expanding its role beyond DNA repair.

    Evidence Reciprocal Co-IP and colocalization with DKC1, in vitro ssRNA glycosylase assay, 47S rRNA RIP, and siRNA depletion with rRNA maturation/5-hmU readouts

    PMID:23246433

    Open questions at the time
    • Direct RNA substrate base in vivo not pinpointed
    • Mechanistic coupling between glycosylase activity and rRNA processing unclear
  13. 2012 Medium

    Linked SMUG1 to replication recovery after 5-FU exposure, distinguishing a recovery-phase role from acute uracil/FU excision.

    Evidence SMUG1/UNG knockdown in human cells, recovery-phase FU sensitivity, cell-cycle and comet assays, and CHK1 phosphorylation immunoblot

    PMID:23253900

    Open questions at the time
    • Single lab, knockdown only
    • Mechanistic link between SMUG1 and CHK1/replication checkpoint not established
  14. 2014 High

    Quantified SMUG1's endogenous contribution to immunoglobulin class switching and A:T hypermutation as the principal residual activity in Ung-/- mice.

    Evidence Ung-/- Smug1-/- double-knockout mice, in vitro switching assays, serum isotype quantification, and SHM pattern analysis

    PMID:24771041

    Open questions at the time
    • Did not define the molecular step at which SMUG1 acts in switching
    • Strand-specific processing at switch regions not resolved
  15. 2017 High

    Defined the kinetic mechanism of lesion recognition, establishing a wedge-based search and product-release-limited catalysis with detectable enzyme and DNA conformational changes.

    Evidence Stopped-flow Trp/2-aminopurine fluorescence and FRET pre-steady-state kinetics on dU:dG DNA

    PMID:29051947

    Open questions at the time
    • Did not identify factors that accelerate product release in cells
    • Recognition kinetics for oxidized substrates not measured
  16. 2017 High

    Established the in vivo lesion-specific consequences of SMUG1 loss, with selective genomic 5-hmU accumulation and synergistic uracil buildup and CpG-biased mutational signatures upon combined UNG loss.

    Evidence Single/double knockout mice, mass-spectrometric genomic lesion quantification, and whole-genome sequencing of tumors

    PMID:28775312

    Open questions at the time
    • Origin of CpG-biased mutations not mechanistically dissected
    • Tissue-specific tumor risk not quantified
  17. 2019 High

    Extended the RNA role to telomerase biogenesis, showing SMUG1 is required for co-transcriptional hTERC processing and DKC1 binding, with telomerase deficiency and impaired bone marrow proliferation on its loss.

    Evidence hTERC RIP, DKC1-hTERC binding in SMUG1-depleted cells, processing-intermediate analysis, telomerase activity assays, and Smug1-knockout mouse bone marrow phenotype

    PMID:31412240

    Open questions at the time
    • Whether catalytic glycosylase activity is required for hTERC processing unresolved
    • Identity of the modified bases regulating hTERC not fully defined
  18. 2023 High

    Identified UV-DDB as a stimulatory partner that relieves SMUG1 product inhibition during 5-hmdU repair, addressing how tight AP-site binding is overcome in cells.

    Evidence In vitro excision assays with purified proteins, EMSA, single-molecule analysis, immunofluorescence colocalization, and proximity ligation assay

    PMID:36971122

    Open questions at the time
    • Whether UV-DDB stimulates SMUG1 on other substrates not tested
    • Generality across cell types and lesion contexts not established

Open questions

Synthesis pass · forward-looking unresolved questions
  • How SMUG1's glycosylase chemistry mechanistically couples to its RNA quality-control roles in rRNA and hTERC processing, and whether catalytic activity is required, remains unresolved.
  • Catalytic-dead separation-of-function not reported in the corpus
  • Direct in vivo RNA base substrate not identified
  • Regulation distinguishing DNA-repair versus RNA-processing functions unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140097 catalytic activity, acting on DNA 7 GO:0016787 hydrolase activity 4 GO:0003677 DNA binding 3 GO:0003723 RNA binding 2 GO:0140098 catalytic activity, acting on RNA 1
Localization
GO:0005634 nucleus 2 GO:0005730 nucleolus 1
Pathway
R-HSA-73894 DNA Repair 5 R-HSA-168256 Immune System 2 R-HSA-8953854 Metabolism of RNA 2
Partners
APE1DDB1DDB2DKC1UNG

Evidence

Reading pass · 19 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2001 SMUG1 was identified as the major uracil-DNA glycosylase activity in UNG-deficient mice, functioning as the primary backup enzyme for repair of deaminated cytosine. SMUG1 is present at similar levels in non-proliferating and proliferating tissues, indicating a replication-independent role in DNA repair, distinct from UNG which localizes to replication foci. Specific neutralizing antibodies used to identify SMUG1 activity in Ung-/- mouse cells and tissues; subcellular localization by cell fractionation The EMBO journal High 11483530
2001 Human SMUG1 (hSMUG1) was identified as identical to HMUDG (5-hydroxymethyluracil DNA N-glycosylase), demonstrating that SMUG1 excises 5-hydroxymethyluracil (5hmUra) from DNA with ~20x the specific activity of the most purified bovine HMUDG fraction. Protein purification from calf thymus, SDS-PAGE renaturation assay, mass spectrometry peptide identification, recombinant GST-fusion protein expression and in vitro enzymatic assay The Journal of biological chemistry High 11526119
2003 Crystal structure of SMUG1 complexed with DNA and base-excision products revealed a more invasive interaction with dsDNA than other UDGs, and an elegant water displacement/replacement mechanism that allows SMUG1 to exclude thymine while accepting 5-hydroxymethyluracil (HmU) as a substrate. SMUG1 is specialized for antimutational uracil excision and also excises the oxidation-damage product HmU. X-ray crystallography of SMUG1-DNA complex Molecular cell High 12820976
2003 Rat and human SMUG1 excise 5-formyluracil (fU), uracil (U), 5-hydroxyuracil (hoU), and 5-hydroxymethyluracil (hmU) from both single-stranded and double-stranded DNA, but not analogous cytosine derivatives (5-hydroxycytosine, 5-formylcytosine) or other oxidized damage. hSMUG1 antibodies neutralized fU, hmU, and hoU activities in HeLa cell extracts, confirming SMUG1 is the primary repair enzyme for this subset of oxidized pyrimidines. Recombinant protein expression of rat and human SMUG1, in vitro substrate specificity assays with various modified bases in ss and dsDNA, antibody neutralization of HeLa cell extract activity Biochemistry High 12718543
2004 Site-directed mutagenesis of hSMUG1 identified crucial residues for catalysis and damage recognition: Asn85 and His239 are required for N-glycosidic bond hydrolysis; Phe98 mediates pi-pi stacking discrimination of pyrimidine rings; Asn163 forms specific hydrogen bonds to the Watson-Crick face of the base; and Gly87-Met91 loop enables recognition of C5 substituents through water-bridged (uracil) or direct (hoU, hmU, fU) hydrogen bonds. Site-directed mutagenesis of hSMUG1 combined with in vitro activity assays for multiple substrates (U, hoU, hmU, fU) and homology modeling based on Xenopus laevis SMUG1 structure Nucleic acids research High 15466595
2005 siRNA-mediated silencing of Smug1 in mouse embryo fibroblasts generates a mutator phenotype with increased C:G to T:A transitions. Cells deficient in both Smug1 and Ung show an additive 10-fold increase in spontaneous C:G to T:A transitions at non-CpG sites, demonstrating that these enzymes have distinct and non-redundant roles. Smug1-deficient cells are also hypersensitive to ionizing radiation, revealing a role of SMUG1 in repair of oxidative cytosine lesions. Stable siRNA-mediated knockdown of Smug1, spontaneous mutation frequency measurement (lacZ reporter), double-mutant epistasis analysis (Ung-/- Smug1 knockdown), ionizing radiation sensitivity assay The EMBO journal High 15902269
2006 SMUG1, if overexpressed, can partially substitute for UNG in antibody diversification (somatic hypermutation and class switch recombination), but SMUG1 plays little natural role in antibody diversification because it is diminishingly expressed during B-cell activation. Even when overexpressed, SMUG1 favors conventional repair of U:G lesions rather than diversification, distinguishing it from UNG which associates with replication sites. SMUG1 overexpression in DT40 B cells and transgenic mice; somatic hypermutation pattern analysis; isotype switching assays in SMUG-transgenic msh2-/- ung-/- mice; expression analysis during B-cell activation The EMBO journal High 16407970
2007 SMUG1 (but not UNG) excises 5-fluorouracil (FU) incorporated into DNA and protects against FU cytotoxicity. Accumulation of FU in the genome, rather than uracil excision, is the predominant cause of FU cytotoxicity in mammalian cells. Gene-targeted cell lines defective in UNG, SMUG1, or both; FU cytotoxicity assays; genomic FU/uracil incorporation measurements Cancer research High 17283124
2007 SMUG1 and UNG2 coordinate the initial steps of base excision repair by distinct mechanisms: UNG2 stimulates AP-site cleavage by APE1 and lacks product-binding capacity enabling rapid turnover; SMUG1 binds tightly to AP-sites and inhibits AP-site cleavage by AP-endonucleases. A specific motif important for AP-site product binding was identified in SMUG1; mutations in this motif increase catalytic turnover by reducing product binding. In vitro AP-site binding and cleavage assays; E. coli complementation assay; site-directed mutagenesis of AP-site binding motif; comparison of UNG2 and SMUG1 enzymatic properties Nucleic acids research High 17537817
2008 SMUG1 (bacterial ortholog from Geobacter metallireducens and human SMUG1) are not only uracil-DNA glycosylases but also xanthine DNA glycosylases (XDGs). Mutational analysis identified M57 and H210 as critical for both XDG and UDG activities via interactions with the C2 carbonyl oxygen; G60Y abolishes both activities by blocking base entry to the binding pocket; G63P switches Gme SMUG1 to an exclusive UDG by altering active-site loop flexibility. Site-directed mutagenesis of bacterial and human SMUG1, in vitro XDG and UDG activity assays on ss and dsDNA substrates, molecular dynamics simulations Journal of molecular biology High 18835277
2012 Targeted inactivation of mouse Smug1 ablates nearly all hmU-DNA excision activity in tissue extracts and renders embryo fibroblasts resistant to 5-hydroxymethyldeoxyuridine toxicity. Combined Smug1/Ung knockout leads to loss of nearly all detectable uracil excision activity, confirming SMUG1 as the dominant glycosylase for hmU-excision and major UNG-backup for U-excision in mice. Knockout mouse generation, biochemical assay of tissue extracts, 5-hydroxymethyldeoxyuridine cytotoxicity assay in embryo fibroblasts, double-knockout epistasis Nucleic acids research High 22447450
2012 SMUG1 is predominantly double-strand-specific under physiological salt conditions (in the presence of Mg2+ and monovalent salts), whereas UNG2 efficiently removes uracil from both ss and dsDNA. Uracil in AID hotspot sequences is removed 200-fold more efficiently from ssDNA by UNG2 than by SMUG1, explaining why SMUG1 cannot replace UNG2 in antibody diversification despite not being excluded from Ig loci. In vitro UDG activity assays under varying ionic conditions with ss and dsDNA substrates; measurement of sequence preference for AID hotspot sequences DNA repair High 22483865
2012 SMUG1 interacts with the pseudouridine synthase Dyskerin (DKC1) and colocalizes with DKC1 in nucleoli and Cajal bodies. SMUG1 has enzymatic activity on single-stranded RNA containing 5-hydroxymethyldeoxyuridine. SMUG1 associates with the 47S rRNA precursor, and depletion of SMUG1 reduces mature rRNA levels with concomitant increase in polyadenylated rRNA and accumulation of 5-hydroxymethyluridine in rRNA. Co-immunoprecipitation (SMUG1-DKC1 interaction), colocalization by immunofluorescence, in vitro RNA glycosylase activity assay, RNA immunoprecipitation (47S rRNA), siRNA depletion with rRNA maturation analysis and 5-hydroxymethyluridine quantification Molecular cell High 23246433
2012 Loss of SMUG1 (but not UNG) leads to >2-fold increased sensitivity to 5-fluorouracil specifically after drug removal (recovery phase), associated with prolonged S-phase arrest, transient increase in DNA double-strand breaks, and altered CHK1 phosphorylation, indicating SMUG1 has a role in resumption of replication following 5-FU treatment. SMUG1 and UNG knockdown in human cell lines, 5-FU sensitivity assays (continuous vs. 24h then recovery), cell cycle analysis, comet assay, CHK1 phosphorylation by immunoblot Mutation research Medium 23253900
2014 Endogenous SMUG1 contributes to immunoglobulin class switching in Ung-/- mice, with most residual class switching in Ung-/- mice depending on SMUG1. In vitro switching to IgG1 and serum IgG3, IgG2b, and IgA are greatly diminished in Ung-/- Smug1-/- mice. SMUG1 deficiency in an Ung-/- background further reduces somatic hypermutation at A:T base pairs. Double-knockout mice (Ung-/- Smug1-/-), in vitro class switching assays, serum immunoglobulin isotype quantification, somatic hypermutation pattern analysis European journal of immunology High 24771041
2017 Pre-steady-state kinetic analysis revealed the mechanism of damaged base recognition by hSMUG1: the enzyme undergoes nonspecific DNA binding followed by specific damaged-base recognition using a 'wedge' strategy for lesion search. SMUG1 is rate-limited by release from the AP-site product (tight product binding confirmed), with the true catalytic rate faster than kcat measured at steady state. Conformational changes in both enzyme (Trp fluorescence) and DNA (2-aminopurine fluorescence and FRET) were detected during recognition. Stopped-flow fluorescence spectroscopy using Trp fluorescence, 2-aminopurine fluorescence, and FRET with dU:dG DNA substrate; pre-steady-state kinetic analysis Molecular bioSystems High 29051947
2017 In Smug1-/- mice, 5-hydroxymethyluracil accumulates in genomic DNA (up to 26-fold in brain, correlating with 5-hydroxymethylcytosine levels), while uracil does not accumulate. Ung-/- Smug1-/- mice show synergistic uracil accumulation (up to 25-fold). Whole genome sequencing of double-deficient tumors reveals C to T transitions primarily at CpG sequences, identifying mutational signatures of combined UNG/SMUG1 deficiency. Knockout mice (single and double), genomic 5-hydroxymethyluracil and uracil quantification by mass spectrometry, whole-genome sequencing of tumors for mutational signature analysis Scientific reports High 28775312
2019 SMUG1 interacts with the telomeric RNA component (hTERC) and is required for co-transcriptional processing of nascent hTERC into mature form. SMUG1 regulates base modifications in hTERC in a region between the CR4/CR5 domain and H box; loss of SMUG1 leads to reduced DKC1 binding to hTERC, accumulation of 3'-polyadenylated and 3'-extended hTERC intermediates degraded by an EXOSC10-independent pathway, and telomerase deficiency leading to impaired bone marrow proliferation in Smug1-knockout mice. RNA immunoprecipitation (SMUG1-hTERC interaction), DKC1-hTERC binding assay in SMUG1-depleted cells, hTERC processing intermediate analysis by northern blot/RT-PCR, telomerase activity assay, Smug1-knockout mouse bone marrow proliferation analysis Cell reports High 31412240
2023 UV-DDB (UV-damaged DNA-binding protein, consisting of DDB1 and DDB2 subunits) stimulates SMUG1 excision activity on 5-hmdU substrates by 4-5-fold in vitro, displaces SMUG1 from abasic site products (EMSA), and decreases SMUG1 half-life on DNA ~8-fold (single-molecule analysis). In cells, DDB2 and SMUG1 form discrete co-localizing foci after 5-hmdU treatment and exhibit transient interaction by proximity ligation assay. In vitro biochemical excision assay with purified proteins, EMSA, single-molecule analysis, immunofluorescence colocalization, proximity ligation assay, siRNA knockdown with poly(ADP-ribose) accumulation readout Nucleic acids research High 36971122

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1998 FDG accumulation and tumor biology. Nuclear medicine and biology 271 9639291
2017 FDG PET for therapy monitoring in Hodgkin and non-Hodgkin lymphomas. European journal of nuclear medicine and molecular imaging 204 28411336
1998 Carbon-11-thymidine and FDG to measure therapy response. Journal of nuclear medicine : official publication, Society of Nuclear Medicine 171 9776283
2001 Excision of deaminated cytosine from the vertebrate genome: role of the SMUG1 uracil-DNA glycosylase. The EMBO journal 165 11483530
2009 123I-MIBG scintigraphy and 18F-FDG PET in neuroblastoma. Journal of nuclear medicine : official publication, Society of Nuclear Medicine 127 19617326
2001 Definitive identification of mammalian 5-hydroxymethyluracil DNA N-glycosylase activity as SMUG1. The Journal of biological chemistry 117 11526119
2007 5-Fluorouracil incorporated into DNA is excised by the Smug1 DNA glycosylase to reduce drug cytotoxicity. Cancer research 114 17283124
2003 Structure and specificity of the vertebrate anti-mutator uracil-DNA glycosylase SMUG1. Molecular cell 112 12820976
2003 Mammalian 5-formyluracil-DNA glycosylase. 2. Role of SMUG1 uracil-DNA glycosylase in repair of 5-formyluracil and other oxidized and deaminated base lesions. Biochemistry 102 12718543
2020 FAPI PET/CT: Will It End the Hegemony of 18F-FDG in Oncology? Journal of nuclear medicine : official publication, Society of Nuclear Medicine 101 33277397
2005 C --> T mutagenesis and gamma-radiation sensitivity due to deficiency in the Smug1 and Ung DNA glycosylases. The EMBO journal 100 15902269
2007 Uracil-DNA glycosylases SMUG1 and UNG2 coordinate the initial steps of base excision repair by distinct mechanisms. Nucleic acids research 95 17537817
2014 Thoracic staging in lung cancer: prospective comparison of 18F-FDG PET/MR imaging and 18F-FDG PET/CT. Journal of nuclear medicine : official publication, Society of Nuclear Medicine 90 24504054
2012 Germline ablation of SMUG1 DNA glycosylase causes loss of 5-hydroxymethyluracil- and UNG-backup uracil-excision activities and increases cancer predisposition of Ung-/-Msh2-/- mice. Nucleic acids research 86 22447450
2014 The Basic Principles of FDG-PET/CT Imaging. PET clinics 81 26050942
2012 New PET radiopharmaceuticals beyond FDG for brain tumor imaging. The quarterly journal of nuclear medicine and molecular imaging : official publication of the Italian Association of Nuclear Medicine (AIMN) [and] the International Association of Radiopharmacology (IAR), [and] Section of the Society of... 79 22617239
2010 Characterization of T-cell lymphomas by FDG PET/CT. AJR. American journal of roentgenology 78 20651187
2006 SMUG1 is able to excise uracil from immunoglobulin genes: insight into mutation versus repair. The EMBO journal 73 16407970
1998 Imaging of mediastinal lymph nodes: CT, MR, and FDG PET. Radiographics : a review publication of the Radiological Society of North America, Inc 73 9747607
2013 FDG-PET imaging in HIV infection and tuberculosis. Seminars in nuclear medicine 69 23905617
2009 Pathophysiologic correlation between 62Cu-ATSM and 18F-FDG in lung cancer. Journal of nuclear medicine : official publication, Society of Nuclear Medicine 68 19910425
2006 Evaluation of NTHL1, NEIL1, NEIL2, MPG, TDG, UNG and SMUG1 genes in familial colorectal cancer predisposition. BMC cancer 68 17029639
2014 Regulation of 18F-FDG accumulation in colorectal cancer cells with mutated KRAS. Journal of nuclear medicine : official publication, Society of Nuclear Medicine 65 25453050
2012 The human base excision repair enzyme SMUG1 directly interacts with DKC1 and contributes to RNA quality control. Molecular cell 62 23246433
2020 FDG PET/CT in the Staging of Lung Cancer. Current radiopharmaceuticals 61 31868151
2014 Oncogene pathway activation in mammary tumors dictates FDG-PET uptake. Cancer research 59 25239452
2010 18F-FPPRGD2 and 18F-FDG PET of response to Abraxane therapy. Journal of nuclear medicine : official publication, Society of Nuclear Medicine 56 21149494
2016 PET Tracers Beyond FDG in Prostate Cancer. Seminars in nuclear medicine 53 27825431
2014 Relationship between 18F-FDG accumulation and lactate dehydrogenase A expression in lung adenocarcinomas. Journal of nuclear medicine : official publication, Society of Nuclear Medicine 53 25342384
2010 The use of FDG-PET to target tumors by radiotherapy. Strahlentherapie und Onkologie : Organ der Deutschen Rontgengesellschaft ... [et al] 50 20814658
2014 FDG-PET and amyloid-PET imaging: the diverging paths. Current opinion in neurology 49 24927239
2017 Uracil Accumulation and Mutagenesis Dominated by Cytosine Deamination in CpG Dinucleotides in Mice Lacking UNG and SMUG1. Scientific reports 47 28775312
2016 FDG-PET imaging in hematological malignancies. Blood reviews 42 27090170
2010 Non-FDG PET in the practice of oncology. Indian journal of cancer 40 20448372
2004 Mutational analysis of the damage-recognition and catalytic mechanism of human SMUG1 DNA glycosylase. Nucleic acids research 40 15466595
2017 Benign Bone Conditions That May Be FDG-avid and Mimic Malignancy. Seminars in nuclear medicine 38 28583274
2014 Uracil excision by endogenous SMUG1 glycosylase promotes efficient Ig class switching and impacts on A:T substitutions during somatic mutation. European journal of immunology 38 24771041
2015 Characterizing POEMS Syndrome with 18F-FDG PET/CT. Journal of nuclear medicine : official publication, Society of Nuclear Medicine 36 26182964
2012 Strikingly different properties of uracil-DNA glycosylases UNG2 and SMUG1 may explain divergent roles in processing of genomic uracil. DNA repair 36 22483865
2011 Lymphomatoid granulomatosis: CT and FDG-PET findings. Korean journal of radiology 35 22043148
1997 Possible role of FDG-PET in the evaluation of urologic malignancies. Anticancer research 34 9179213
2006 Cancer screening with FDG-PET. The quarterly journal of nuclear medicine and molecular imaging : official publication of the Italian Association of Nuclear Medicine (AIMN) [and] the International Association of Radiopharmacology (IAR), [and] Section of the Society of... 33 16557201
2022 Systematically evaluating DOTATATE and FDG as PET immuno-imaging tracers of cardiovascular inflammation. Scientific reports 32 35418569
2013 Single-strand selective monofunctional uracil-DNA glycosylase (SMUG1) deficiency is linked to aggressive breast cancer and predicts response to adjuvant therapy. Breast cancer research and treatment 32 24253812
2019 FDG PET-CT in pediatric Langerhans cell histiocytosis. Pediatric blood & cancer 31 31599488
2016 FDG-PET characteristics of Hürthle cell and follicular adenomas. Annals of nuclear medicine 31 27221817
2022 Clozapine induces astrocyte-dependent FDG-PET hypometabolism. European journal of nuclear medicine and molecular imaging 30 35122511
2019 Obligatory role of endoplasmic reticulum in brain FDG uptake. European journal of nuclear medicine and molecular imaging 29 30617965
2010 Role of FDG PET-CT in recurrent renal cell carcinoma. Nuclear medicine communications 29 20661166
2021 PET/CT in Multiple Myeloma: Beyond FDG. Frontiers in oncology 28 33569348
2019 SMUG1 Promotes Telomere Maintenance through Telomerase RNA Processing. Cell reports 28 31412240
2015 High resolution FDG-microPET of carotid atherosclerosis: plaque components underlying enhanced FDG uptake. The international journal of cardiovascular imaging 28 26280889
2021 FDG-PET in presymptomatic C9orf72 mutation carriers. NeuroImage. Clinical 27 34049163
2022 Signaling Pathways That Drive 18F-FDG Accumulation in Cancer. Journal of nuclear medicine : official publication, Society of Nuclear Medicine 26 35241480
2021 The VISION Forward: Recognition and Implication of PSMA-/18F-FDG+ mCRPC. Journal of nuclear medicine : official publication, Society of Nuclear Medicine 26 34933889
2009 18F-FDG PET/CT of transitional cell carcinoma. AJR. American journal of roentgenology 26 19933624
2008 Insights from xanthine and uracil DNA glycosylase activities of bacterial and human SMUG1: switching SMUG1 to UDG. Journal of molecular biology 26 18835277
2022 Dual-Tracer PET/CT Protocol with [18F]-FDG and [68Ga]Ga-FAPI-46 for Cancer Imaging: A Proof of Concept. Journal of nuclear medicine : official publication, Society of Nuclear Medicine 25 35422446
2012 FDG for therapy of metabolically active tumors. Seminars in nuclear medicine 24 22475427
2012 Low-sensitivity FDG-PET studies: less common lung neoplasms. Seminars in nuclear medicine 24 22681674
2003 Assessment of therapy response by FDG PET in pediatric patients. The quarterly journal of nuclear medicine : official publication of the Italian Association of Nuclear Medicine (AIMN) [and] the International Association of Radiopharmacology (IAR) 24 12714953
2019 18F-FDG PET/CT IN DIFFERENTIATED THYROID CARCINOMA. Acta endocrinologica (Bucharest, Romania : 2005) 23 31508177
2017 Pre-steady-state kinetic analysis of damage recognition by human single-strand selective monofunctional uracil-DNA glycosylase SMUG1. Molecular bioSystems 23 29051947
2017 Interim FDG-PET Imaging in Lymphoma. Seminars in nuclear medicine 23 29195613
2017 Gut microbiota and physiologic bowel 18F-FDG uptake. EJNMMI research 22 28861740
2019 Genetic features associated with 18F-FDG uptake in intrahepatic cholangiocarcinoma. Annals of surgical treatment and research 21 30941318
2023 From FDG and beyond: the evolving potential of nuclear medicine. Annals of nuclear medicine 20 37749301
2015 FDG-PET/CT findings in systemic mastocytosis: a French multicentre study. European journal of nuclear medicine and molecular imaging 20 26140850
2012 SMUG1 but not UNG DNA glycosylase contributes to the cellular response to recovery from 5-fluorouracil induced replication stress. Mutation research 20 23253900
2015 Multiple myeloma: 18F-FDG-PET/CT and diagnostic imaging. Seminars in nuclear medicine 18 25475376
2007 Sensitivity of FDG PET, GLUT1 expression and proliferative index in bronchioloalveolar lung cancer. Nuclear medicine communications 18 17264775
2022 Interim FDG-PET/CT for Response Assessment of Lymphoma. Seminars in nuclear medicine 17 36376131
2022 FDG-PET in Autoimmune Encephalitis: Utility, Pattern of Abnormalities, and Correlation with Autoantibodies. Annals of Indian Academy of Neurology 17 36911487
2016 FDG-PET study of patients with Leigh syndrome. Journal of the neurological sciences 17 26944169
2016 The correlation between FDG uptake and biological molecular markers in pancreatic cancer patients. European journal of radiology 17 27666620
2014 FDG-PET imaging for Hodgkin lymphoma: current use and future applications. Clinical advances in hematology & oncology : H&O 17 25000313
2010 FDG-PET/CT in pediatric solid tumors. The quarterly journal of nuclear medicine and molecular imaging : official publication of the Italian Association of Nuclear Medicine (AIMN) [and] the International Association of Radiopharmacology (IAR), [and] Section of the Society of... 17 20823808
2021 The Multiple Cellular Roles of SMUG1 in Genome Maintenance and Cancer. International journal of molecular sciences 16 33671338
2021 Hypermetabolic pulmonary lesions on FDG-PET/CT: Tularemia or neoplasia? Infectious diseases now 16 34242840
2018 Heterogeneous brain FDG-PET metabolic patterns in patients with C9orf72 mutation. Neurological sciences : official journal of the Italian Neurological Society and of the Italian Society of Clinical Neurophysiology 16 30554355
2017 FDG-PET Scan: a new Paradigm for Follicular Lymphoma Management. Mediterranean journal of hematology and infectious diseases 16 28512558
2020 Association between immunotherapy biomarkers and glucose metabolism from F-18 FDG PET. European review for medical and pharmacological sciences 15 32894535
2014 Reproducibility of FDG PET based metabolic tumor volume measurements and of their FDG distribution within. The quarterly journal of nuclear medicine and molecular imaging : official publication of the Italian Association of Nuclear Medicine (AIMN) [and] the International Association of Radiopharmacology (IAR), [and] Section of the Society of... 15 24695005
2008 Limitations of FDG-PET and FDG-PET with computed tomography for detecting synchronous cancer in pharyngeal cancer. Archives of otolaryngology--head & neck surgery 15 19015450
2021 RNA Metabolism Guided by RNA Modifications: The Role of SMUG1 in rRNA Quality Control. Biomolecules 14 33430019
2017 Science to Practice: Does FDG Differentiate Morphologically Unstable from Stable Atherosclerotic Plaque? Radiology 14 28318446
2005 FDG-PET in the management of germ cell tumor. Annals of oncology : official journal of the European Society for Medical Oncology 14 15923438
2004 FDG-PET scanning in the diagnosis of gastrointestinal cancers. Scandinavian journal of gastroenterology. Supplement 14 15696855
2021 Clinical and FDG-PET/CT correlates in patients with polymyalgia rheumatica. Clinical and experimental rheumatology 13 33822711
2019 Interest of FDG-PET in the Management of Mantle Cell Lymphoma. Frontiers in medicine 13 31024918
2015 FDG-PET/CT in abdominal post-transplant lymphoproliferative disease. The British journal of radiology 13 26544161
2015 Association Between (18)F-FDG Avidity and the BRAF Mutation in Papillary Thyroid Carcinoma. Nuclear medicine and molecular imaging 13 26941858
2024 Predictive [18F]-FDG PET/CT-Based Radiogenomics Modelling of Driver Gene Mutations in Non-small Cell Lung Cancer. Academic radiology 12 38997880
2022 FDG-PET findings associated with various medical procedures and treatments. Japanese journal of radiology 12 36575286
2020 Nivolumab-Induced Periaortitis Demonstrated by FDG PET/CT. Clinical nuclear medicine 12 32701815
2018 The use of F-FDG PET/CT in testicular cancer. Translational andrology and urology 12 30456190
2016 Monitoring Vasculitis with 18F-FDG PET. The quarterly journal of nuclear medicine and molecular imaging : official publication of the Italian Association of Nuclear Medicine (AIMN) [and] the International Association of Radiopharmacology (IAR), [and] Section of the Society of... 12 27280732
2023 UV-DDB stimulates the activity of SMUG1 during base excision repair of 5-hydroxymethyl-2'-deoxyuridine moieties. Nucleic acids research 11 36971122
2022 A Role of Non-FDG Tracers in Lung Cancer? Seminars in nuclear medicine 11 35803770
2012 FDG PET and FDG PET/CT in patients with gastrointestinal stromal tumours. Wiener medizinische Wochenschrift (1946) 11 22890522

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