Affinage

GEN1

Flap endonuclease GEN homolog 1 · UniProt Q17RS7

Length
908 aa
Mass
102.9 kDa
Annotated
2026-06-10
100 papers in source corpus 20 papers cited in narrative 20 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

GEN1 is a structure-specific endonuclease of the Rad2/XPG family that resolves Holliday junctions (HJs) to maintain genome stability during recombinational DNA repair (PMID:20634321). Although largely monomeric in solution, GEN1 dimerizes specifically on the HJ to assemble two symmetrically aligned active sites, and resolves the junction by a coordinated nick-and-counter-nick mechanism in which a rate-limiting first incision triggers a rapid second incision in the continuous strands, yielding ligatable nicked duplexes (PMID:20634321, PMID:26578604); single-molecule kinetics show the monomer binds and distorts the junction first, then remains bound until dimerization completes forward resolution, with higher affinity for the singly-cleaved intermediate enforcing productive cleavage (PMID:30590761). Crystal structures reveal an elaborated FEN-XPG fold adapted for four-way junction recognition, including a chromodomain that directly contacts DNA and is required for catalysis, and capture how cleavage relaxes the junction center to accelerate the second incision (PMID:26686639, PMID:26682650). GEN1 acts on physiological single and double HJs and under-replicated DNA, and it is one of several redundant HJ-processing activities in human cells, operating alongside the SLX4-associated nucleases MUS81-EME1 and SLX1—co-loss of GEN1 and SLX4 is synthetically lethal through failed mitotic resolution of unprocessed junctions (PMID:24080495, PMID:28049850). GEN1 is spatially restricted by a nuclear export signal that confines its activity to mitosis when the nuclear envelope breaks down, temporally separating it from the S-phase BTR (BLM) pathway and preventing inappropriate crossover formation (PMID:25209024, PMID:24831703). At mitosis GEN1 cleaves under-replicated DNA at common fragile sites to enable mitotic DNA synthesis and proper chromosome segregation (PMID:36729836). Independent of its nuclease role, GEN1 promotes kidney development: it interacts with SIX1 to enhance SIX1/EYA1 transcriptional activity and supports GDNF/RET signaling during ureteric bud branching, and its disruption in mice causes CAKUT phenotypes (PMID:29483821). GEN1 activity is also controlled by ubiquitin-mediated turnover via a CUL4-DDB1/WDR-23 pathway (PMID:31409866).

Mechanistic history

Synthesis pass · year-by-year structured walk · 14 steps
  1. 2010 High

    Established the core biochemical identity of GEN1 as a Holliday junction resolvase, answering how a Rad2/XPG-family flap endonuclease could symmetrically cleave a four-way junction.

    Evidence In vitro reconstitution, EM, and nuclease/dimerization assays with human GEN1; HJ resolvase and separation-of-function genetics for the C. elegans ortholog GEN-1

    PMID:20634321 PMID:20661466

    Open questions at the time
    • Atomic basis of junction recognition and dimerization not yet defined
    • Cellular timing and regulation of resolvase activity unknown
  2. 2013 High

    Placed GEN1 within a redundant cellular HJ-resolution network, showing it is one of multiple nucleases (with SLX4-MUS81-EME1/SLX1) required to clear replication-induced junctions.

    Evidence RNAi knockdown of GEN1 and SLX4 in human cells with synthetic-lethality and mitotic phenotype analysis

    PMID:24080495

    Open questions at the time
    • Did not define when each nuclease acts in the cell cycle
    • Mechanism of synthetic lethality at the DNA level not resolved
  3. 2014 High

    Defined the spatial and temporal logic of GEN1 action, showing a nuclear export signal restricts it to mitosis and prevents competition with non-crossover repair pathways.

    Evidence NES mutagenesis and nuclear-localized GEN1 variants with imaging and crossover assays; siRNA depletion with cell-cycle staging and segregation phenotypes

    PMID:24831703 PMID:25209024

    Open questions at the time
    • Whether additional regulation supplements NES-based control in humans unclear
    • How mitotic access is coordinated with envelope breakdown not mechanistically detailed
  4. 2014 High

    Confirmed conserved homodimeric resolvase mechanism and revealed kinetic asymmetry favoring productive resolution, using a thermostable ortholog amenable to biochemistry.

    Evidence Purification and kinetic nuclease assays of Chaetomium thermophilum GEN1 on four-way junctions

    PMID:25315822

    Open questions at the time
    • Ortholog dimerizes constitutively, unlike human GEN1's HJ-induced dimerization
    • Structural basis of accelerated second-strand cleavage not yet shown
  5. 2015 High

    Provided the structural mechanism: crystal structures revealed an elaborated FEN-XPG fold and a catalytically required chromodomain, and explained how cleavage relaxes the junction to accelerate the counter-nick.

    Evidence X-ray crystallography of eukaryotic and human GEN1-DNA complexes with permanganate/2-AP probing, chromodomain truncations, and yeast complementation; full-length human GEN1 nick/counter-nick dissection with phosphorothioate substrates

    PMID:26578604 PMID:26682650 PMID:26686639

    Open questions at the time
    • Structure of the catalytically active dimer on an intact junction not captured
    • Role of the chromodomain in vivo not fully tested
  6. 2017 High

    Extended GEN1 activity to physiological substrates, showing it cleaves large single and double HJs within recombination intermediates with only weak sequence bias.

    Evidence In vitro reconstitution of physiological recombination intermediates with sequence-preference analysis

    PMID:28049850

    Open questions at the time
    • In vivo substrate spectrum and frequency of use versus other nucleases not quantified
  7. 2019 High

    Resolved the single-molecule choreography of resolution, explaining how monomer-to-dimer transitions and intermediate-state affinities enforce committed, forward HJ cleavage.

    Evidence Single-molecule FRET imaging and kinetic analysis of human GEN1-HJ binding and cleavage

    PMID:30590761

    Open questions at the time
    • Dimerization partner stoichiometry in chromatin context unknown
    • How cellular regulation intersects with these kinetics not addressed
  8. 2018 High

    Demonstrated conserved phosphorylation-based suppression of resolvase activity in meiosis, showing the yeast ortholog Yen1 must be held inactive for proper crossover patterning.

    Evidence Phosphorylation-refractory Yen1 mutants with genome-wide crossover mapping and MutLgamma epistasis

    PMID:29920281

    Open questions at the time
    • Human GEN1 is regulated by localization rather than phosphorylation, so direct transfer of mechanism is uncertain
    • Meiotic role of mammalian GEN1 not directly established here
  9. 2023 High

    Identified a specific mitotic genome-maintenance role, showing GEN1 cleaves under-replicated DNA at common fragile sites to enable MiDAS and faithful segregation.

    Evidence GEN1 knockout cells with CFS expression, MiDAS, anaphase bridge, and DNA-damage readouts

    PMID:36729836

    Open questions at the time
    • Whether GEN1 acts directly on fragile-site structures or via HJ intermediates not distinguished
    • Coordination with MUS81 at CFS not fully defined
  10. 2012 Medium

    Suggested a non-canonical centrosomal role, with GEN1 targeting and a catalysis-independent function in preventing centrosome amplification.

    Evidence Immunofluorescence localization, N-terminal signal mapping, and shRNA knockdown with catalytic-dead and localization mutants

    PMID:23166748

    Open questions at the time
    • Single-lab finding not independently confirmed
    • Tension with later evidence that GEN1 is nuclear-excluded/mitosis-restricted unresolved
    • Molecular basis of centrosomal function unknown
  11. 2017 Medium

    Showed GEN1 activity must be restrained at stressed replication forks, with SLX4 physically preventing GEN1-dependent unscheduled DSBs.

    Evidence SLX4-mutant and FA-P patient cells, RuvA ectopic expression, and GEN1 chromatin-fractionation/DSB assays

    PMID:28290553

    Open questions at the time
    • Single-lab study
    • How SLX4 mechanistically blocks GEN1 chromatin association not defined
  12. 2016 Medium

    Established in vivo functional redundancy between Gen1 and Eme1 in mammalian DNA repair and meiosis through mouse genetics.

    Evidence Gen1/Eme1 double-mutant mice, MEF damage-sensitivity assays, and meiotic recombination analysis

    PMID:27383418

    Open questions at the time
    • Single-lab study
    • Molecular substrates underlying redundancy not directly identified
  13. 2019 Medium

    Identified ubiquitin-mediated regulation of resolvase abundance, placing the C. elegans ortholog under CUL4-DDB1/WDR-23 control to tune DSB repair.

    Evidence C. elegans WDR-23 isoform-specific mutants, epistasis, DSB repair assays, and proteasome inhibition

    PMID:31409866

    Open questions at the time
    • Conservation of this regulation for human GEN1 not tested
    • Direct ubiquitination of GEN1 not demonstrated
  14. 2020 Medium

    Defined a developmental, recombination-independent function: GEN1 interacts with SIX1 and drives GDNF/RET signaling to control ureteric bud branching and kidney morphogenesis.

    Evidence Gen1 knockout/mutant mice, Co-IP with SIX1, transcriptional reporters, RNA-seq, ex vivo kidney culture, and double-mutant analysis with Robo2

    PMID:29483821 PMID:32226308 PMID:35071283

    Open questions at the time
    • Single research group
    • Whether the developmental role requires GEN1 nuclease activity not established
    • Direct mechanism by which GEN1 enhances SIX1/EYA1 transcription unclear

Open questions

Synthesis pass · forward-looking unresolved questions
  • How GEN1's catalytic genome-maintenance function is mechanistically reconciled with its centrosomal and developmental (SIX1/GDNF) roles, and whether these are nuclease-dependent, remains unresolved.
  • No structure of the active dimer on an intact junction
  • Human-specific ubiquitin regulation untested
  • Mechanistic link between resolvase activity and transcriptional/centrosomal functions undefined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003677 DNA binding 3 GO:0016787 hydrolase activity 3 GO:0140097 catalytic activity, acting on DNA 3 GO:0140110 transcription regulator activity 1
Localization
GO:0005634 nucleus 2 GO:0005815 microtubule organizing center 1 GO:0005829 cytosol 1
Pathway
R-HSA-1266738 Developmental Biology 3 R-HSA-1640170 Cell Cycle 3 R-HSA-73894 DNA Repair 3
Partners
EME1EYA1MUS81SIX1SLX1SLX4

Evidence

Reading pass · 20 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2010 GEN1 is a monomeric 5'-flap endonuclease of the Rad2/XPG family that binds specifically to Holliday junctions (HJs) and resolves them by a dual incision mechanism introducing symmetrically related nicks in the pair of continuous (noncrossing) strands within the lifetime of the GEN1-HJ complex. Uniquely among Rad2/XPG family members, GEN1 dimerizes on HJs, providing two symmetrically aligned active sites required for resolution. Biochemical reconstitution in vitro, electron microscopy (structural characterization of GEN1-HJ complex), nuclease activity assays, dimerization analysis Genes & development High 20634321
2010 The C. elegans ortholog GEN-1 has Holliday junction resolvase biochemical activity and facilitates repair of DNA double-strand breaks in vivo. Its DNA damage-signaling function (promoting germ cell cycle arrest and apoptosis via a pathway parallel to the canonical RPA/CHK1/CEP-1 pathway) is separable from its DNA repair role. GEN-1 acts redundantly with the 9-1-1 complex for genome stability. Forward genetic screen, biochemical activity assays, mutational analysis, epistasis analysis in C. elegans PLoS genetics High 20661466
2013 GEN1 and SLX4-associated nucleases (MUS81-EME1 and SLX1) are each essential for resolution of replication-induced Holliday junctions in human cells. Depletion of both SLX4 and GEN1 is synthetically lethal due to dysfunctional mitosis in the presence of unprocessed HJs. In vivo HJ resolution requires both SLX4-associated MUS81-EME1 and SLX1 acting in concert. RNAi knockdown of GEN1 and SLX4 in human cells, synthetic lethality analysis, cell viability and mitosis phenotype assays Cell reports High 24080495
2014 GEN1 is regulated exclusively by nuclear exclusion via a nuclear export signal (NES), restricting its activity to mitosis when the nuclear membrane breaks down (independent of phosphorylation, unlike yeast Yen1). A nuclear-localized GEN1 partially suppresses phenotypes of BLM/MUS81 loss but causes elevated crossover formation, demonstrating that spatial control of GEN1 promotes genome stability by preventing competition with non-crossover promoting repair pathways. NES identification and mutagenesis, construction of nuclear-localized GEN1 variant, live-cell imaging, crossover frequency assays, epistasis with BLM and MUS81 Nature communications High 25209024
2014 Cells lacking SLX-MUS and GEN1 exhibit chromosome missegregation, micronucleus formation, elevated 53BP1-positive G1 nuclear bodies, impaired replication fork movement, S-phase progression defects, endogenous checkpoint activation, chromosome segmentation, and multinucleation, demonstrating that GEN1 and the SLX-MUS complex act at temporally distinct phases of the cell cycle (GEN1 at mitosis) for HJ processing, while cycling cells preferentially use the BTR complex in S phase. siRNA depletion of GEN1, SLX4, and MUS81 in human cells; cell cycle analysis; chromosome segregation assays; replication fork analysis; 53BP1 foci quantification Genes & development High 24831703
2014 GEN1 from thermophilic fungus Chaetomium thermophilum (CtGEN1) cleaves four-way DNA junctions as a discrete homodimer with nanomolar affinity. Second-strand cleavage occurs an order of magnitude faster than first-strand cleavage, ensuring productive resolution. These properties closely resemble bacterial HJ resolvases. Protein expression and purification, in vitro nuclease assay, cruciform cleavage kinetics, gel electrophoresis, binding affinity measurements Journal of molecular biology High 25315822
2015 Crystal structure of eukaryotic GEN1 bound to DNA at 2.5 Å reveals an elaborated FEN-XPG family fold modified for four-way junction resolution. The functional unit is a GEN1 monomer bound to the resolution cleavage product; within the crystal lattice a GEN1 dimer interface juxtaposes two products that can be reconnected into a four-way junction. Cleavage is accompanied by a relaxation/opening of the DNA structure at the center, explaining how second-strand cleavage is accelerated. X-ray crystallography (2.5 Å resolution), permanganate probing, 2-aminopurine fluorescence Cell reports High 26686639
2015 Crystal structure of human GEN1 complexed with DNA at 3.0 Å reveals that GEN1 contains a chromodomain (not previously found in nucleases) that directly contacts DNA and is required for catalytic activity. Truncation of the chromodomain severely impairs GEN1 cleavage activity. Structure-guided mutations validated mechanistic findings in vitro and in yeast. X-ray crystallography (3.0 Å), chromodomain truncation mutants, in vitro nuclease assays, yeast complementation eLife High 26682650
2015 Full-length human GEN1 promotes HJ resolution by a nick and counter-nick mechanism with dual coordinated incisions producing ligatable nicked duplex products. GEN1 is largely monomeric in solution but dimerizes on the HJ; first nick occurs upon dimerization. First-strand cleavage is rate limiting; second-strand cleavage is rapid. The two incisions can be uncoupled using phosphorothioate-containing non-cleavable linkages. Purification of full-length human GEN1, in vitro HJ cleavage assays, phosphorothioate substrate analysis, native gel electrophoresis, dimerization assays Nucleic acids research High 26578604
2017 GEN1 efficiently cleaves both single and double Holliday junctions contained within large physiological recombination intermediates, not just synthetic DNA substrates. GEN1 exhibits a weak sequence preference for incision between two G residues in a T-rich region, distinct from the strict consensus required by E. coli RuvC. In vitro reconstitution of recombination intermediates, nuclease activity assays, sequence preference analysis Proceedings of the National Academy of Sciences of the United States of America High 28049850
2018 CDK-mediated phosphorylation of yeast Yen1 (GEN1 ortholog) prevents its recruitment to recombination intermediates during meiotic prophase I. Yen1 phosphorylation-refractory mutants resolve DNA joint molecules prematurely, form crossovers independently of MutLγ, and impair spatial distribution of crossover events genome-wide, demonstrating that active suppression of Yen1/GEN1 activity is required for meiotic crossover patterning. Phosphorylation-refractory yeast mutant construction, meiotic recombination analysis, genome-wide crossover mapping, epistasis with MutLγ Developmental cell High 29920281
2019 Single-molecule FRET shows that human GEN1 monomer initially binds the HJ following its isomer bias and distorts it for catalysis. GEN1 monomer remains tightly bound without dissociation until a GEN1 dimer forms and the HJ is fully resolved. Fast on-rates and slow off-rates of GEN1 dimer, and increased affinity for singly-cleaved HJ, enforce forward resolution. GEN1 monomer binds singly-cleaved HJ more tightly than intact HJ, providing a fail-safe mechanism. Single-molecule FRET imaging, kinetic analysis of GEN1-HJ binding and cleavage Nucleic acids research High 30590761
2012 GEN1 is a centrosome-associated protein with an N-terminal centrosome localization signal sufficient for centrosome targeting. GEN1 depletion causes aberrant centrosome numbers, multiple spindle poles, multinucleation, increased apoptosis, elevated spontaneous DNA damage, and severely impaired homologous recombination. Centrosome association (but not catalytic activity) of GEN1 is required to prevent centrosome hyper-amplification and multiple mitotic spindles. Immunofluorescence localization, N-terminal signal mapping and truncation, shRNA knockdown with phenotypic readouts, GEN1 complementation with catalytic-dead and localization mutants PloS one Medium 23166748
2017 SLX4 prevents unscheduled GEN1-dependent DSBs at demised replication forks during replication stress. This function is independent of SLX4's interaction with endonucleases and requires the physical presence of SLX4. Ectopic expression of HJ-binding protein RuvA inhibits DSBs in SLX4-deficient cells by blocking GEN1 chromatin-association, rescuing genome integrity. RNAi and FA-P patient cells with SLX4 mutants, RuvA ectopic expression, chromatin fractionation to detect GEN1 association, DSB quantification Scientific reports Medium 28290553
2016 Gen1 and Eme1 (MUS81-EME1 complex component) play redundant roles in DNA repair and meiotic recombination in mice. Combined homozygous mutations of Gen1 and Eme1 are synthetically lethal during early embryogenesis. Gen1 mutations alone do not cause DNA repair deficiency in MEFs but sensitize heterozygous Eme1 mutant MEFs to DNA-damaging agents and reduce meiotic recombination efficiency in Eme1 mutant mice. Mouse genetics (homozygous Gen1 and Eme1 double mutants), MEF DNA damage sensitivity assays, meiotic recombination analysis DNA and cell biology Medium 27383418
2018 Disruption of Gen1 in mice leads to renal agenesis, duplex kidney, hydronephrosis, and vesicoureteral reflux (CAKUT phenotypes). GEN1 interacts with SIX1 and enhances transcriptional activity of the SIX1/EYA1 complex, a key regulator of the GDNF morphogen. Gen1 mutation impairs Grem1 and Gdnf expression, resulting in excessive ureteric bud formation and defective ureteric bud branching during early kidney development. Gen1 knockout mouse model, co-immunoprecipitation of GEN1 with SIX1, transcriptional reporter assays, in situ hybridization for Grem1/Gdnf, ureteric bud phenotype analysis International journal of biological sciences Medium 29483821
2019 In C. elegans, GEN-1 (Holliday junction resolvase) is an evolutionarily conserved substrate of the CUL4-DDB1 E3 ubiquitin ligase receptor WDR-23. Nuclear WDR-23B isoform inhibits GEN-1 activity, most likely by promoting protein turnover via ubiquitination, while cytoplasmic WDR-23A performs a proteasome-independent role. Differential ubiquitination by distinct WDR-23 isoforms modulates double-strand break repair. C. elegans WDR-23 isoform-specific mutants, epistasis analysis, DSB repair assays, proteasome inhibition experiments Scientific reports Medium 31409866
2023 GEN1 promotes common fragile site (CFS) expression by cleaving under-replicated DNA at CFS loci, enabling mitotic DNA synthesis (MiDAS). Loss of GEN1 reduces CFS expression, causing defects in MiDAS, reduced ultrafine anaphase bridge formation, increased bichromatid constrictions (consistent with unresolved under-replicated DNA), and DNA damage in the ensuing cell cycle due to aberrant chromosome segregation. GEN1 knockout cells, CFS expression assays, MiDAS quantification, anaphase bridge and bichromatid constriction analysis, DNA damage marker quantification Cell reports High 36729836
2020 Gen1 mutation in mice impairs late metanephric development: decreases ureteric bud branching (confirmed by ex vivo kidney primordia culture), causes kidney hypoplasia, and impairs programmed apoptosis at the end of the nephric duct, leading to abnormal ureter-bladder connections (VUR or UVJO). RNA-seq analysis reveals Gen1 mutation impairs expression of multiple metanephric regulatory genes including Six2. Mouse Gen1 mutant model, ex vivo kidney culture, RNA-seq transcriptome analysis, ureteric bud branching quantification International journal of biological sciences Medium 32226308
2022 Robo2 and Gen1 act synergistically in mouse kidney development to regulate ureteric bud formation: double disruption of Robo2 and Gen1 significantly increases CAKUT phenotypes (especially duplicated collecting system) with increased ectopic UB formation. The two genes promote cell proliferation by activating the GDNF/RET pathway and downstream MAPK/ERK signaling. Double-mutant mouse model (piggyBac transposon), embryonic ureteric bud analysis, GDNF/RET and MAPK/ERK pathway readouts Frontiers in medicine Medium 35071283

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2016 The natural history of cutaneous propionibacteria, and reclassification of selected species within the genus Propionibacterium to the proposed novel genera Acidipropionibacterium gen. nov., Cutibacterium gen. nov. and Pseudopropionibacterium gen. nov. International journal of systematic and evolutionary microbiology 314 27488827
2020 A phylogenomic and comparative genomic framework for resolving the polyphyly of the genus Bacillus: Proposal for six new genera of Bacillus species, Peribacillus gen. nov., Cytobacillus gen. nov., Mesobacillus gen. nov., Neobacillus gen. nov., Metabacillus gen. nov. and Alkalihalobacillus gen. nov. International journal of systematic and evolutionary microbiology 206 31617837
2015 Review of the taxonomy of the genus Arthrobacter, emendation of the genus Arthrobacter sensu lato, proposal to reclassify selected species of the genus Arthrobacter in the novel genera Glutamicibacter gen. nov., Paeniglutamicibacter gen. nov., Pseudoglutamicibacter gen. nov., Paenarthrobacter gen. nov. and Pseudarthrobacter gen. nov., and emended description of Arthrobacter roseus. International journal of systematic and evolutionary microbiology 181 26486726
2014 Characterization of Romboutsia ilealis gen. nov., sp. nov., isolated from the gastro-intestinal tract of a rat, and proposal for the reclassification of five closely related members of the genus Clostridium into the genera Romboutsia gen. nov., Intestinibacter gen. nov., Terrisporobacter gen. nov. and Asaccharospora gen. nov. International journal of systematic and evolutionary microbiology 177 24480908
2010 Mechanism of Holliday junction resolution by the human GEN1 protein. Genes & development 130 20634321
2013 Human GEN1 and the SLX4-associated nucleases MUS81 and SLX1 are essential for the resolution of replication-induced Holliday junctions. Cell reports 122 24080495
2014 Roles of SLX1-SLX4, MUS81-EME1, and GEN1 in avoiding genome instability and mitotic catastrophe. Genes & development 116 24831703
1999 The family Coriobacteriaceae: reclassification of Eubacterium exiguum (Poco et al. 1996) and Peptostreptococcus heliotrinreducens (Lanigan 1976) as Slackia exigua gen. nov., comb. nov. and Slackia heliotrinireducens gen. nov., comb. nov., and Eubacterium lentum (Prevot 1938) as Eggerthella lenta gen. nov., comb. nov. International journal of systematic bacteriology 114 10319481
1999 Description of Cellulophaga baltica gen. nov., sp. nov. and Cellulophaga fucicola gen. nov., sp. nov. and reclassification of [Cytophaga] lytica to Cellulophaga lytica gen. nov., comb. nov. International journal of systematic bacteriology 101 10425785
2013 Fusicatenibacter saccharivorans gen. nov., sp. nov., isolated from human faeces. International journal of systematic and evolutionary microbiology 97 23625266
2014 Spatial control of the GEN1 Holliday junction resolvase ensures genome stability. Nature communications 80 25209024
2010 The Caenorhabditis elegans homolog of Gen1/Yen1 resolvases links DNA damage signaling to DNA double-strand break repair. PLoS genetics 79 20661466
2000 Reassessment of the taxonomic structure of the diazotrophic genus Azoarcus sensu lato and description of three new genera and new species, Azovibrio restrictus gen. nov., sp. nov., Azospira oryzae gen. nov., sp. nov. and Azonexus fungiphilus gen. nov., sp. nov. International journal of systematic and evolutionary microbiology 77 10758872
2010 GEN1/Yen1 and the SLX4 complex: Solutions to the problem of Holliday junction resolution. Genes & development 76 20203129
2014 Roseimaritima ulvae gen. nov., sp. nov. and Rubripirellula obstinata gen. nov., sp. nov. two novel planctomycetes isolated from the epiphytic community of macroalgae. Systematic and applied microbiology 58 25487949
2000 Description of Mogibacterium pumilum gen. nov., sp. nov. and Mogibacterium vescum gen. nov., sp. nov., and reclassification of Eubacterium timidum (Holdeman et al. 1980) as Mogibacterium timidum gen. nov., comb. nov. International journal of systematic and evolutionary microbiology 56 10758875
2011 A revision of Cyanonectria and Geejayessia gen. nov., and related species with Fusarium-like anamorphs. Studies in mycology 47 21523191
2012 Description of Sphingorhabdus planktonica gen. nov., sp. nov. and reclassification of three related members of the genus Sphingopyxis in the genus Sphingorhabdus gen. nov. International journal of systematic and evolutionary microbiology 46 22798658
2006 Hatena arenicola gen. et sp. nov., a katablepharid undergoing probable plastid acquisition. Protist 45 16891155
2014 Pantanalinema gen. nov. and Alkalinema gen. nov.: novel pseudanabaenacean genera (Cyanobacteria) isolated from saline-alkaline lakes. International journal of systematic and evolutionary microbiology 44 25351877
2022 Establishment of Genome Based Criteria for Classification of the Family Desulfovibrionaceae and Proposal of Two Novel Genera, Alkalidesulfovibrio gen. nov. and Salidesulfovibrio gen. nov. Frontiers in microbiology 42 35694308
2019 Next-Gen Approaches to Flavor-Related Metabolism. Annual review of plant biology 42 30835532
2013 The mystery of clade X: Orciraptor gen. nov. and Viridiraptor gen. nov. are highly specialised, algivorous amoeboflagellates (Glissomonadida, Cercozoa). Protist 40 23994723
2010 Halobellus clavatus gen. nov., sp. nov. and Halorientalis regularis gen. nov., sp. nov., two new members of the family Halobacteriaceae. International journal of systematic and evolutionary microbiology 40 21169458
2015 Crystal Structure of a Eukaryotic GEN1 Resolving Enzyme Bound to DNA. Cell reports 37 26686639
2014 Viewing protein fitness landscapes through a next-gen lens. Genetics 35 25316787
2000 Actinoalloteichus cyanogriseus gen. nov., sp. nov. International journal of systematic and evolutionary microbiology 35 10939647
2017 Resolution of single and double Holliday junction recombination intermediates by GEN1. Proceedings of the National Academy of Sciences of the United States of America 34 28049850
2015 GEN1 promotes Holliday junction resolution by a coordinated nick and counter-nick mechanism. Nucleic acids research 34 26578604
2008 Proteiniborus ethanoligenes gen. nov., sp. nov., an anaerobic protein-utilizing bacterium. International journal of systematic and evolutionary microbiology 33 18175674
2016 Cutaneous hyalohyphomycosis due to Parengyodontium album gen. et comb. nov. Medical mycology 32 27161787
2015 Human Holliday junction resolvase GEN1 uses a chromodomain for efficient DNA recognition and cleavage. eLife 31 26682650
2007 Sediminibacter furfurosus gen. nov., sp. nov. and Gilvibacter sediminis gen. nov., sp. nov., novel members of the family Flavobacteriaceae. International journal of systematic and evolutionary microbiology 30 17267962
2015 Hold your horSSEs: controlling structure-selective endonucleases MUS81 and Yen1/GEN1. Frontiers in genetics 29 26284109
2018 Regulated Crossing-Over Requires Inactivation of Yen1/GEN1 Resolvase during Meiotic Prophase I. Developmental cell 28 29920281
2005 Dokdonia donghaensis gen. nov., sp. nov., isolated from sea water. International journal of systematic and evolutionary microbiology 28 16280490
2020 Anaerohalosphaera lusitana gen. nov., sp. nov., and Limihaloglobus sulfuriphilus gen. nov., sp. nov., isolated from solar saltern sediments, and proposal of Anaerohalosphaeraceae fam. nov. within the order Sedimentisphaerales. International journal of systematic and evolutionary microbiology 25 31804915
2016 Crusticorallina gen. nov., a nongeniculate genus in the subfamily Corallinoideae (Corallinales, Rhodophyta). Journal of phycology 25 27434825
2012 miRNA data analysis: next-gen sequencing. Methods in molecular biology (Clifton, N.J.) 25 22144206
2015 Rhabdobacter roseus gen. nov., sp. nov., isolated from soil. International journal of systematic and evolutionary microbiology 24 26508300
2014 Methyloparacoccus murrellii gen. nov., sp. nov., a methanotroph isolated from pond water. International journal of systematic and evolutionary microbiology 24 24676728
2008 Alkanibacter difficilis gen. nov., sp. nov. and Singularimonas variicoloris gen. nov., sp. nov., hexane-degrading bacteria isolated from a hexane-treated biofilter. International journal of systematic and evolutionary microbiology 21 18842850
2007 Methylohalomonas lacus gen. nov., sp. nov. and Methylonatrum kenyense gen. nov., sp. nov., methylotrophic gammaproteobacteria from hypersaline lakes. International journal of systematic and evolutionary microbiology 21 18048722
2018 Rubneribacter badeniensis gen. nov., sp. nov. and Enteroscipio rubneri gen. nov., sp. nov., new members of the Eggerthellaceae isolated from human faeces. International journal of systematic and evolutionary microbiology 20 29537365
2022 Taxogenomic status of phylogenetically distant Frankia clusters warrants their elevation to the rank of genus: A description of Protofrankia gen. nov., Parafrankia gen. nov., and Pseudofrankia gen. nov. as three novel genera within the family Frankiaceae. Frontiers in microbiology 19 36425027
2018 Molecular and Morphological Diversity of the Oxymonad Genera Monocercomonoides and Blattamonas gen. nov. Protist 19 30138782
2016 Harryflintia acetispora gen. nov., sp. nov., isolated from chicken caecum. International journal of systematic and evolutionary microbiology 19 27432404
2012 A novel role of human holliday junction resolvase GEN1 in the maintenance of centrosome integrity. PloS one 19 23166748
2019 Nuclear and cytoplasmic WDR-23 isoforms mediate differential effects on GEN-1 and SKN-1 substrates. Scientific reports 18 31409866
2015 Taxonomic revision of Pseudolachnea and Pseudolachnella and establishment of Neopseudolachnella and Pseudodinemasporium gen. nov. Mycologia 18 25572100
2015 Reclassification of Actinobacillus muris as Muribacter muris gen. nov., comb. nov. International journal of systematic and evolutionary microbiology 18 26296776
2015 Chujaibacter soli gen. nov., sp. nov., isolated from soil. Journal of microbiology (Seoul, Korea) 18 26310302
2018 GEN-1 immunotherapy for the treatment of ovarian cancer. Future oncology (London, England) 17 30325199
2014 GEN1 from a thermophilic fungus is functionally closely similar to non-eukaryotic junction-resolving enzymes. Journal of molecular biology 17 25315822
2025 Precision Metal Nanoclusters Meet Proteins: Crafting Next-Gen Hybrid Materials. ACS nano 16 39854644
2023 Phylogenomic and molecular markers based studies on Staphylococcaceae and Gemella species. Proposals for an emended family Staphylococcaceae and three new families (Abyssicoccaceae fam. nov., Salinicoccaceae fam. nov. and Gemellaceae fam. nov.) harboring four new genera, Lacicoccus gen. nov., Macrococcoides gen. nov., Gemelliphila gen. nov., and Phocicoccus gen. nov. Antonie van Leeuwenhoek 16 37523090
2017 Halodesulfovibrio spirochaetisodalis gen. nov. sp. nov. and reclassification of four Desulfovibrio spp. International journal of systematic and evolutionary microbiology 16 27902290
2010 Mutation and association analysis of GEN1 in breast cancer susceptibility. Breast cancer research and treatment 16 20512659
2007 Reclassification of Aquaspirillum itersonii and Aquaspirillum peregrinum as Novispirillum itersonii gen. nov., comb. nov. and Insolitispirillum peregrinum gen. nov., comb. nov. International journal of systematic and evolutionary microbiology 16 18048733
2023 Testudinimyces gracilis gen. nov, sp. nov. and Astrotestudinimyces divisus gen. nov, sp. nov., two novel, deep-branching anaerobic gut fungal genera from tortoise faeces. International journal of systematic and evolutionary microbiology 15 37252853
2019 Resolution of the Holliday junction recombination intermediate by human GEN1 at the single-molecule level. Nucleic acids research 15 30590761
2018 Disruption of Gen1 Causes Congenital Anomalies of the Kidney and Urinary Tract in Mice. International journal of biological sciences 15 29483821
2010 Fibrisoma limi gen. nov., sp. nov., a filamentous bacterium isolated from tidal flats. International journal of systematic and evolutionary microbiology 15 20601484
2021 Neglectibacter timonensis gen. nov., sp. nov. and Scatolibacter rhodanostii gen. nov., sp. nov., two anaerobic bacteria isolated from human stool samples. Archives of microbiology 14 34932147
2015 Pontivivens insulae gen. nov., sp. nov., isolated from seawater. International journal of systematic and evolutionary microbiology 13 26012580
2014 Paradevosia shaoguanensis gen. nov., sp. nov., isolated from a coking wastewater. Current microbiology 13 25234654
2023 Amygdalobacter indicium gen. nov., sp. nov., and Amygdalobacter nucleatus sp. nov., gen. nov.: novel bacteria from the family Oscillospiraceae isolated from the female genital tract. International journal of systematic and evolutionary microbiology 12 37787404
2016 Gen1 and Eme1 Play Redundant Roles in DNA Repair and Meiotic Recombination in Mice. DNA and cell biology 12 27383418
2014 Halopolyspora alba gen. nov., sp. nov., isolated from sediment. International journal of systematic and evolutionary microbiology 12 24860112
2024 Novel thermophilic genera Geochorda gen. nov. and Carboxydochorda gen. nov. from the deep terrestrial subsurface reveal the ecophysiological diversity in the class Limnochordia. Frontiers in microbiology 11 39376703
2021 Reclassification of Facklamia ignava, Facklamia sourekii and Facklamia tabacinasalis as Falseniella ignava gen. nov., comb. nov., Hutsoniella sourekii gen. nov., comb. nov., and Ruoffia tabacinasalis gen. nov., comb. nov., and description of Ruoffia halotolerans sp. nov., isolated from hypersaline Inland Sea of Qatar. Antonie van Leeuwenhoek 11 34181136
2017 SLX4 Prevents GEN1-Dependent DSBs During DNA Replication Arrest Under Pathological Conditions in Human Cells. Scientific reports 11 28290553
2017 Chthonobacter albigriseus gen. nov., sp. nov., isolated from grass-field soil. International journal of systematic and evolutionary microbiology 11 28475028
2016 Compostibacter hankyongensis gen. nov.,  sp. nov., isolated from compost. International journal of systematic and evolutionary microbiology 11 27335062
2015 Youhaiella tibetensis gen. nov., sp. nov., isolated from subsurface sediment. International journal of systematic and evolutionary microbiology 11 25829329
2015 Parasediminibacterium paludis gen. nov., sp. nov., isolated from wetland. International journal of systematic and evolutionary microbiology 11 26514371
2012 Neoscardovia arbecensis gen. nov., sp. nov., isolated from porcine slurries. Systematic and applied microbiology 11 22824582
2024 Circular RNA vaccines: Pioneering the next-gen cancer immunotherapy. Cancer pathogenesis and therapy 10 40697518
2023 GEN1 promotes common fragile site expression. Cell reports 10 36729836
2021 Tenebriella gen. nov. - The dark twin of Oscillatoria. Molecular phylogenetics and evolution 10 34391914
2020 Gen1 mutation caused kidney hypoplasia and defective ureter-bladder connections in mice. International journal of biological sciences 10 32226308
2017 Agarilytica rhodophyticola gen. nov., sp. nov., isolated from Gracilaria blodgettii. International journal of systematic and evolutionary microbiology 10 28879838
2016 Enterovirga rhinocerotis gen. nov., sp. nov., isolated from Rhinoceros unicornis faeces. Antonie van Leeuwenhoek 10 28032205
2014 The persistent contributions of RNA to eukaryotic gen(om)e architecture and cellular function. Cold Spring Harbor perspectives in biology 10 25081515
2012 Barrientosiimonas humi gen. nov., sp. nov., an actinobacterium of the family Dermacoccaceae. International journal of systematic and evolutionary microbiology 10 22389286
2025 Phycobium rhodophyticola gen. nov., sp. nov. and Aliiphycobium algicola gen. nov., sp. nov., isolated from the phycosphere of marine red algae. Journal of microbiology (Seoul, Korea) 9 40598999
2020 Descriptions of Roseiconus nitratireducens gen. nov. sp. nov. and Roseiconus lacunae sp. nov. Archives of microbiology 9 33047175
2018 Xylanibacillus composti gen. nov., sp. nov., isolated from compost. International journal of systematic and evolutionary microbiology 9 29458465
2016 Effect of GEN1 interference on the chemosensitivity of the breast cancer MCF-7 and SKBR3 cell lines. Oncology letters 9 27284361
2015 Pseudohalocynthiibacter aestuariivivens gen. nov., sp. nov., isolated from a tidal flat. International journal of systematic and evolutionary microbiology 9 25713042
2023 Phylogenomic analysis of the genus Alcanivorax: proposal for division of this genus into the emended genus Alcanivorax and two novel genera Alloalcanivorax gen. nov. and Isoalcanivorax gen. nov. International journal of systematic and evolutionary microbiology 8 36748586
2021 Vescimonas gen. nov., Vescimonas coprocola sp. nov., Vescimonas fastidiosa sp. nov., Pusillimonas gen. nov. and Pusillimonas faecalis sp. nov. isolated from human faeces. International journal of systematic and evolutionary microbiology 8 34726590
2020 Seongchinamella unica gen. nov., sp. nov., isolated from a tidal mudflat of beach, and transfer of Halioglobus sediminis to Seongchinamella sediminis comb. nov. and Halioglobus lutimaris to Pseudohalioglobus gen. nov. as Pseudohalioglobus lutimaris comb. nov. International journal of systematic and evolutionary microbiology 8 32100687
2018 Oceanibium sediminis gen. nov., sp. nov., isolated from marine sediment. International journal of systematic and evolutionary microbiology 8 30484760
2017 Antarcticibacterium flavum gen. nov., sp. nov., isolated from marine sediment. International journal of systematic and evolutionary microbiology 8 29205128
2025 Toward Sustainable Polyhydroxyalkanoates: A Next-Gen Biotechnology Approach. Polymers 7 40219244
2023 Salinilacihabitans rarus gen. nov., sp. nov., Natrononativus amylolyticus gen. nov., sp. nov., Natronobeatus ordinarius gen. nov., sp. nov., and Halovivax gelatinilyticus sp. nov., halophilic archaea, isolated from a salt lake and soda lakes. Extremophiles : life under extreme conditions 7 37400737
2022 Robo2 and Gen1 Coregulate Ureteric Budding by Activating the MAPK/ERK Signaling Pathway in Mice. Frontiers in medicine 7 35071283
2022 Huaxiibacter chinensis gen. nov., sp. nov., recovered from human sputum. International journal of systematic and evolutionary microbiology 7 35976100
2021 Sandaracinobacteroides hominis gen. nov., sp. nov., isolated from human skin. Archives of microbiology 7 34302507

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