Affinage

ORC2

Mitochondrial ornithine transporter 2 · UniProt Q9BXI2

Length
301 aa
Mass
32.6 kDa
Annotated
2026-06-10
31 papers in source corpus 21 papers cited in narrative 20 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

ORC2 is a conserved subunit of the origin recognition complex that is essential for the initiation of chromosomal DNA replication, binding chromatin prior to DNA synthesis and prior to loading of replication licensing factors (PMID:8552193, PMID:8552194). Within human ORC, ORC2 provides an assembly platform: a single ORC assembly domain mediates interaction with all other ORC subunits and two nuclear localization signals drive nuclear accumulation, with ORC2 existing as ORC(2-5) subcomplexes that require ORC1 to load onto chromatin (PMID:16762929), and co-expression with ORC1 suppresses ORC1-induced apoptosis (PMID:16537645). Its winged-helix domain binds double-stranded DNA through a flexible β-hairpin anchor (R540, K548, K549) using a topology distinct from archaeal and yeast ORC (PMID:30963726). ORC2 chromatin association is dynamically controlled across the cell cycle by post-translational modification: cyclin/CDK phosphorylation at Thr116 and Thr226 dissociates ORC2–5 from origins during S phase, and PP1, recruited via the KSVSF motif on ORC2, reverses this in late M phase to permit re-loading (PMID:22334659, PMID:24732362, PMID:24792176, PMID:11900493); Plk1 phosphorylation at Ser188 sustains replication and the pre-replicative complex under replication stress (PMID:21947279, PMID:23188630). At G2/M, PIAS4-mediated SUMOylation by SUMO2 (reversed by SENP2) recruits the histone demethylase KDM5A to centromeres to convert H3K4me3 to H3K4me2, supporting α-satellite transcription, pericentric heterochromatin silencing, and prevention of DNA re-replication (PMID:27052177, PMID:29050267). ORC2 localizes to centrosomes, centromeres, and heterochromatin and associates with HP1, with its loss causing chromosome condensation, congression, and centrosome defects (PMID:15215892), and it regulates replication timing of euchromatin versus heterochromatin (PMID:11137005). Beyond canonical licensing, DNA-bound ORC2 independent of the full six-subunit complex compacts chromatin, attracts or excludes repressive histone marks, and restricts CTCF binding to shape chromosome loops at thousands of genomic sites (PMID:40504688); notably, selected cancer cells survive complete ORC2/ORC5 loss with intact MCM2-7 loading, indicating the intact six-subunit ORC is not strictly required for origin firing in this context (PMID:32989049).

Mechanistic history

Synthesis pass · year-by-year structured walk · 14 steps
  1. 1996 High

    Established that ORC2 is required for the initiation step of chromosomal DNA replication and acts upstream of licensing, defining its core biological role.

    Evidence Immunodepletion of XORC2 from Xenopus egg extracts and chromatin-binding assays; reciprocal genetic and Co-IP analysis of fission yeast Orp2 with Cdc2 and Cdc18

    PMID:8552193 PMID:8552194

    Open questions at the time
    • Did not resolve the molecular mechanism of origin selection
    • Checkpoint signaling role inferred from loss-of-function but not mechanistically dissected
  2. 2000 High

    Showed that ORC2 controls genome-wide replication timing, coupling origin function to chromatin state and mitotic condensation.

    Evidence Genetic mutant analysis with BrdU timing and cytology of mitotic chromosomes in Drosophila larvae

    PMID:11137005

    Open questions at the time
    • Mechanism linking ORC2 to late-replication delay not defined
    • Connection between replication timing and condensation defect correlative
  3. 2004 High

    Defined the cell-cycle-dependent localization of human ORC2 to centrosomes, centromeres, and heterochromatin and its physical association with HP1, expanding its role beyond replication into chromosome architecture.

    Evidence siRNA depletion, immunofluorescence, cell fractionation, and Co-IP with HP1α/HP1β in human cells

    PMID:15215892

    Open questions at the time
    • Whether HP1 association is direct not established
    • Mechanistic basis of centrosome localization unknown
  4. 2006 High

    Mapped the ORC2 assembly domain and NLS signals and showed ORC1-dependence of chromatin loading, clarifying how ORC2 nucleates and positions the complex.

    Evidence Domain-deletion mutant expression, chromatin fractionation, and Co-IP in HeLa cells; ORC1/ORC2 co-expression apoptosis rescue assays

    PMID:16537645 PMID:16762929

    Open questions at the time
    • Structural basis of subunit contacts not resolved
    • ORC1-induced apoptosis pathway only phenomenologically described
  5. 2012 High

    Identified CDK phosphorylation at Thr116/Thr226 as the switch that dissociates ORC2 and partner subunits from origins during S phase, providing a mechanism for replication licensing control.

    Evidence In vitro CDK kinase assay, phosphomimetic/phosphodefective mutagenesis, chromatin fractionation, and origin ChIP

    PMID:22334659

    Open questions at the time
    • Did not identify the re-loading mechanism (resolved later by PP1 work)
  6. 2014 Medium

    Showed PP1 directly binds ORC2 via the KSVSF motif and dephosphorylates Thr116/Thr226 in late M phase, closing the loop on cyclic ORC2 chromatin re-loading.

    Evidence Co-IP of PP1 with ORC2, PP1 inhibitor and isoform manipulation, chromatin fractionation

    PMID:24732362 PMID:24792176

    Open questions at the time
    • PP1 isoform specificity in vivo not fully resolved
    • Single lab across two papers
  7. 2012 High

    Established a stress-responsive arm in which Plk1 phosphorylates ORC2 at Ser188 to maintain the pre-RC and replication under genotoxic stress.

    Evidence In vitro kinase assay, mass-spec phosphosite mapping, S188A mutagenesis, BrdU and checkpoint assays; drug sensitivity and xenograft validation

    PMID:21947279 PMID:23188630

    Open questions at the time
    • Downstream effectors of S188 phosphorylation not defined
    • How Plk1 and CDK phosphorylation are coordinated unclear
  8. 2012 Medium

    Connected ORC2 to stability of the ORC-associated factor ORCA/LRWD1, showing ORC2 binds and protects the non-ubiquitinated form from proteasomal degradation.

    Evidence In vivo ubiquitination assay, reciprocal Co-IP, siRNA knockdown, proteasome inhibition

    PMID:22935713

    Open questions at the time
    • Whether protection is direct steric blocking not shown
    • Single lab
  9. 2016 High

    Revealed a SUMO2-dependent function of ORC2 at G2/M that recruits KDM5A to centromeres, linking ORC2 modification to centromeric histone methylation, α-satellite transcription, and re-replication suppression.

    Evidence In vivo SUMOylation site mapping, Co-IP of KDM5A with SUMO2-ORC2, SUMO-less rescue, chromatin analysis and flow cytometry

    PMID:27052177

    Open questions at the time
    • Direct SUMO-KDM5A interaction interface not defined
    • Whether this is replication-independent not fully separated
  10. 2017 Medium

    Identified the writer/eraser pair (PIAS4/SENP2) governing reversible ORC2 SUMOylation, establishing the enzymatic control of the centromeric methylation circuit.

    Evidence PIAS4 knockdown, SENP2 overexpression, ORC2-SUMO2 fusion rescue, flow cytometry and histone methylation analysis

    PMID:29050267

    Open questions at the time
    • Upstream signals triggering G2/M SUMOylation unknown
    • Single lab extending prior work
  11. 2019 High

    Provided the structural basis for ORC2 DNA engagement, defining a winged-helix domain with a distinct dsDNA-binding topology.

    Evidence Crystal structure, NMR chemical shift perturbation, mutagenesis of R540/K548/K549, molecular dynamics

    PMID:30963726

    Open questions at the time
    • Structure of full ORC2 within the assembled complex not resolved
    • Sequence specificity of DNA binding not addressed
  12. 2020 Medium

    Demonstrated that the intact six-subunit ORC, including ORC2, is dispensable for MCM2-7 loading and origin firing in selected cancer cells, challenging the obligatory role of ORC2 in licensing.

    Evidence CRISPR-Cas9 ORC2/ORC5 knockout, chromatin fractionation, origin firing assays, Western blot in HCT116

    PMID:32989049

    Open questions at the time
    • How origins are loaded without ORC unresolved
    • May reflect adaptation specific to these selected cells
  13. 2024 Medium

    Placed ORC2 origin localization downstream of the ORFIUS (BRD1/HBO1/BRCA1/BARD1) complex, linking ORC2 recruitment to a chromatin-modifying complex and replication stress responsiveness.

    Evidence siRNA/CRISPR depletion of complex components, ORC2 foci immunofluorescence, origin firing assays, ChIP

    PMID:38288445

    Open questions at the time
    • Direct versus indirect recruitment by ORFIUS not separated
    • Mechanism of stress unresponsiveness in BRCA1-mutant cells unclear
  14. 2025 Medium

    Defined a replication-independent genomic role for DNA-bound ORC2 in chromatin compaction, histone mark deposition/protection, and CTCF/loop regulation at thousands of sites.

    Evidence Multi-omics (ChIP-seq, ATAC-seq, Hi-C) in ORC-knockout cancer cell lines

    PMID:40504688

    Open questions at the time
    • No in vitro reconstitution of ORC2-driven compaction
    • Mechanism by which individual subunits bind without co-occupancy unknown

Open questions

Synthesis pass · forward-looking unresolved questions
  • How the multiple ORC2 modification circuits (CDK, PP1, Plk1, SUMO) are integrated, and how replication-independent chromatin functions relate to the dispensability of intact ORC in some cells, remain unresolved.
  • No unified model coordinating the four modification axes
  • Mechanism of ORC-independent MCM loading unknown
  • In vitro reconstitution of the chromatin-architecture role lacking

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003677 DNA binding 3
Localization
GO:0000228 nuclear chromosome 3 GO:0005634 nucleus 2 GO:0005815 microtubule organizing center 1
Pathway
R-HSA-4839726 Chromatin organization 3 R-HSA-69306 DNA Replication 3 R-HSA-1640170 Cell Cycle 2
Complex memberships
Origin recognition complex (ORC)

Evidence

Reading pass · 20 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1996 Xenopus Orc2-related protein (XORC2) is required for chromosomal DNA replication: immunodepletion of XORC2 from Xenopus egg extracts abolishes replication of chromosomal DNA but not elongation synthesis on single-stranded DNA templates. XORC2 binds chromatin prior to DNA synthesis and prior to loading of replication licensing factors. Immunodepletion from Xenopus egg extracts; indirect immunofluorescence Nature High 8552193
1996 Fission yeast Orp2 (ORC2 ortholog) physically interacts with Cdc2 kinase and with the replication activator Cdc18, and is required for DNA replication; cells lacking Orp2 undergo aberrant mitosis indicating Orp2 is involved in generating a checkpoint signal. Genetic interaction screen, co-immunoprecipitation, loss-of-function analysis in S. pombe Nature High 8552194
2000 Drosophila ORC2 is required for normal replication timing: two alleles of Drosophila ORC2 disrupt the normal early-euchromatin/late-heterochromatin replication pattern, causing some euchromatic regions to replicate abnormally late, and resulting in defective mitotic chromosome condensation in those late-replicating regions. Genetic mutant analysis, BrdU incorporation timing assay, cytological examination of mitotic chromosomes in Drosophila larvae Current biology : CB High 11137005
2004 Human ORC2 localizes to centrosomes throughout the entire cell cycle and to centromeres and heterochromatin in a cell-cycle-dependent manner. ORC2 is tightly bound to heterochromatin and HP1α/HP1β during G1 and early S phase, but during late S, G2, and M phases chromatin association is restricted to centromeres. Depletion of ORC2 by siRNA disrupts HP1 localization (without affecting H3K9 methylation), causes S-phase defects with reduced PCNA on chromatin (though MCM proteins remain), induces abnormal chromosome condensation, failed chromosome congression, and multiple centrosomes. siRNA depletion, immunofluorescence, cell fractionation, co-immunoprecipitation with HP1α and HP1β The EMBO journal High 15215892
2006 Human ORC2 contains a single ORC assembly domain required in vivo for interaction with all other ORC subunits, and two nuclear localization signals (NLS) required for ORC accumulation in the nucleus. In the nucleus, ORC2 exists as ORC(2-5) or ORC(1-5) complexes; only ORC(1-5) is chromatin-bound, indicating Orc1 is required to load ORC(2-5) onto chromatin. Additionally, ORC2 suppresses expression of endogenous ORC2, indicating cells limit intracellular ORC2 levels. Stable expression of epitope-tagged domain-deletion mutants in HeLa cells, chromatin fractionation, co-immunoprecipitation The Journal of biological chemistry High 16762929
2006 Co-expression of ORC2 with ORC1 prevents ORC1-induced apoptosis and restores uniform nuclear localization of ORC1. In the absence of ORC2 co-expression, unmodified ORC1 accumulates perinuclearly and rapidly induces p53-independent apoptosis. Transient expression of ORC1 ± ORC2 in cell lines, immunofluorescence, apoptosis assays Journal of cell science Medium 16537645
2011 Polo-like kinase 1 (Plk1) phosphorylates ORC2 at Ser188 in vitro and in vivo. This phosphorylation is enhanced under DNA replication stress (UV, hydroxyurea, gemcitabine, aphidicolin). Cells expressing the unphosphorylatable S188A ORC2 mutant have defective DNA synthesis under stress, fail to maintain functional pre-replicative complex, and activate the intra-S-phase checkpoint. In vitro kinase assay, in vivo phosphorylation (mass spectrometry), site-directed mutagenesis (S188A), BrdU incorporation, pre-RC chromatin binding assays, checkpoint activation assays Molecular and cellular biology High 21947279
2012 Cyclin-dependent kinase phosphorylates ORC2 at Thr-116 and Thr-226 during S phase, causing dissociation of ORC2, ORC3, ORC4, and ORC5 subunits from chromatin and replication origins. Phosphomimetic ORC2 shows defective binding to replication origins and chromatin; phosphodefective ORC2 persists in chromatin binding throughout the cell cycle. In vitro CDK phosphorylation assay, phosphomimetic and phosphodefective mutagenesis, chromatin fractionation, ChIP at replication origins The Journal of biological chemistry High 22334659
2012 ORC2 protects the ORC-associated protein ORCA/LRWD1 from ubiquitin-mediated degradation. ORCA is polyubiquitinated via K48-linked chains by Cul4A-DDB1 E3 ligase at the WD40 repeat domain. ORC2 binds exclusively the non-ubiquitinated form of ORCA, and depletion of ORC2 leads to proteasome-mediated destabilization of ORCA. In vivo ubiquitination assay, co-immunoprecipitation, siRNA knockdown, proteasome inhibitor treatment Cell cycle (Georgetown, Tex.) Medium 22935713
2012 Plk1 phosphorylation of ORC2 (at Ser188) maintains DNA replication under gemcitabine treatment; cells expressing a Plk1-unphosphorylatable ORC2 mutant are more sensitive to gemcitabine than wild-type ORC2-expressing cells. Phosphomimetic/phosphodefective ORC2 mutant expression in cancer cells, drug sensitivity assays, mouse xenograft model Molecular cancer therapeutics Medium 23188630
2014 Protein phosphatase 1 (PP1) physically interacts with ORC2 via the consensus PP1-binding motif 119-KSVSF-123 on ORC2. PP1 dephosphorylates ORC2 at Thr116 and Thr226 in a cell-cycle-dependent manner (late M phase), which is required for re-binding of ORC2 and associated subunits to chromatin and replication origins. Co-immunoprecipitation of PP1 and ORC2, PP1 inhibitor treatment, overexpression and siRNA knockdown of PP1 isoforms, chromatin fractionation Biochemical and biophysical research communications / Biochemical and biophysical research communications Medium 24732362 24792176
2016 ORC2 is SUMOylated by SUMO2 (but not SUMO1) at the G2/M phase of the cell cycle. SUMO2-modified ORC2 recruits the histone demethylase KDM5A to centromeres to convert H3K4me3 to H3K4me2, a permissive histone mark for α-satellite transcription. Loss of ORC2 SUMOylation (SUMO-less ORC2) reduces α-satellite transcription, impairs pericentric heterochromatin silencing, leads to heterochromatin DNA re-replication, activates DNA damage response, and causes polyploidy. In vivo SUMOylation assays, co-immunoprecipitation of KDM5A with SUMO2-ORC2, stable expression of SUMO-less ORC2 mutant, chromatin analysis, flow cytometry Cell reports High 27052177
2016 Papillomavirus E2 protein binds ORC2; however, ORC2 is not detected at the viral origin. ORC2 depletion enhances PV replication and increases E1/E2 occupancy at the viral origin, indicating ORC2 suppresses E2 replicative function rather than promoting viral replication. Over-expression of HPV E2 decreases ORC2 occupation at mammalian replication origins. Co-immunoprecipitation of E2 and ORC2, siRNA depletion of ORC2, transient replication assay, ChIP at viral and cellular origins PLoS pathogens Medium 27701460
2017 ORC2 SUMOylation is reversibly regulated: SUMO E3 ligase PIAS4 promotes ORC2 SUMOylation at G2/M, while de-SUMOylase SENP2 removes it. Depletion of PIAS4 or overexpression of SENP2 eliminates ORC2 SUMOylation, causes abnormal centromeric H3K4 methylation, and results in mitotic bypass and polyploidy; co-expression of ORC2-SUMO2 fusion protein reduces polyploid cell formation. PIAS4 siRNA depletion, SENP2 overexpression, ORC2-SUMO2 fusion rescue, flow cytometry, histone methylation analysis Oncotarget Medium 29050267
2019 The ORC2 winged-helix domain (WHD) binds dsDNA through a flexible β-sheet hairpin anchor region with key residues R540, K548, and K549. NMR chemical shift perturbations reveal a unique dsDNA binding topology distinct from archaeal and yeast ORC WHDs; mutagenesis of these residues validates their importance for binding. Crystal structure determination, NMR backbone assignments and chemical shift perturbation, site-directed mutagenesis of DNA-binding residues, molecular dynamics simulation The FEBS journal High 30963726
2020 Human HCT116 cancer cells can survive and maintain normal MCM2-7 chromatin loading and origin firing even when both ORC2 and ORC5 proteins are eliminated by CRISPR-Cas9 mutation, causing destabilization of ORC1, ORC3, and ORC4 as well. This demonstrates that in these selected cancer cells, the six-subunit ORC is not strictly required for MCM2-7 loading or origin specification. CRISPR-Cas9 gene editing, chromatin fractionation, origin firing assays, Western blot The Journal of biological chemistry Medium 32989049
2024 The ORFIUS complex (BRD1, HBO1, BRCA1, BARD1) promotes ORC2 localization at replication origins. Depletion of BRD1 and/or HBO1 reduces origin firing and reduces the number of nuclei with ORC2 foci. BRCA1 regulates BRD1, HBO1, and ORC2 localization at origins; in BRCA1-mutant HGSC cells, ORC2 remains at origins and is unresponsive to replication stress signals. siRNA/CRISPR depletion of complex components, immunofluorescence for ORC2 foci, origin firing assays, ChIP NAR cancer Medium 38288445
2025 DNA-bound ORC2 (independent of the full six-subunit ORC) compacts chromatin and attracts repressive histone marks at focal genomic sites, while also activating chromatin and protecting genes from repressive marks at other sites. ORC2 also prevents CTCF acquisition at focal sites to regulate chromatin loops and indirectly affects epigenetics. Individual ORC subunits bind thousands of sites without co-occupancy of other subunits. Multi-omics analysis (ChIP-seq, ATAC-seq, Hi-C) in ORC-mutant cancer cell lines (ORC1, ORC2, ORC5 CRISPR knockouts) Cell reports Medium 40504688
2002 Elevated cyclin A-dependent kinase activity in Xenopus egg extract prevents XORC2 from binding to chromatin from permeable erythrocyte nuclei; kinase inhibition reverses this effect. However, inhibiting nuclear-accumulated kinase activity within intact erythrocyte nuclei does not facilitate XORC2 binding to chromatin, suggesting additional mechanisms prevent ORC association within intact terminally differentiated nuclei. Xenopus egg extract replication system, permeable and intact erythrocyte nuclei, cyclin A-CDK titration, kinase inhibitor treatment Experimental cell research Medium 11900493
2011 Sp1 transcription factor participates in recruiting ORC2 to the chicken lysozyme GAS41 replication origin; knockdown of Sp1 by RNA interference reduces specific ORC2 binding to this origin, which maps to a region containing multiple Sp1/Sp3-binding sites coinciding with DNase I hypersensitive sites. Chromatin immunoprecipitation (ChIP), quantitative real-time PCR, siRNA knockdown of Sp1 DNA and cell biology Low 21879882

Source papers

Stage 0 corpus · 31 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2004 Human Orc2 localizes to centrosomes, centromeres and heterochromatin during chromosome inheritance. The EMBO journal 228 15215892
1996 Role for a Xenopus Orc2-related protein in controlling DNA replication. Nature 185 8552193
1996 Interaction of Cdc2 and Cdc18 with a fission yeast ORC2-like protein. Nature 123 8552194
2000 Aberrant replication timing induces defective chromosome condensation in Drosophila ORC2 mutants. Current biology : CB 99 11137005
2012 Plk1 phosphorylation of orc2 and hbo1 contributes to gemcitabine resistance in pancreatic cancer. Molecular cancer therapeutics 70 23188630
2011 Plk1 phosphorylation of Orc2 promotes DNA replication under conditions of stress. Molecular and cellular biology 63 21947279
2006 Ubiquitylation, phosphorylation and Orc2 modulate the subcellular location of Orc1 and prevent it from inducing apoptosis. Journal of cell science 44 16537645
1996 Mouse and human homologues of the yeast origin of replication recognition complex subunit ORC2 and chromosomal localization of the cognate human gene ORC2L. Genomics 41 8808289
2012 Phosphorylation of ORC2 protein dissociates origin recognition complex from chromatin and replication origins. The Journal of biological chemistry 34 22334659
2016 SUMOylated ORC2 Recruits a Histone Demethylase to Regulate Centromeric Histone Modification and Genomic Stability. Cell reports 33 27052177
2019 Circ_ORC2 enhances the regulatory effect of miR-19a on its target gene PTEN to affect osteosarcoma cell growth. Biochemical and biophysical research communications 30 31103262
2020 A human cancer cell line initiates DNA replication normally in the absence of ORC5 and ORC2 proteins. The Journal of biological chemistry 29 32989049
2016 The Replicative Consequences of Papillomavirus E2 Protein Binding to the Origin Replication Factor ORC2. PLoS pathogens 22 27701460
2012 Orc2 protects ORCA from ubiquitin-mediated degradation. Cell cycle (Georgetown, Tex.) 19 22935713
2005 Transcriptional regulation of the Drosophila orc2 gene by the DREF pathway. Biochimica et biophysica acta 17 16343659
2009 Cul4 and DDB1 regulate Orc2 localization, BrdU incorporation and Dup stability during gene amplification in Drosophila follicle cells. Journal of cell science 16 19531585
2006 Genetic analysis of human Orc2 reveals specific domains that are required in vivo for assembly and nuclear localization of the origin recognition complex. The Journal of biological chemistry 15 16762929
2012 Unique pattern of ORC2 and MCM7 localization during DNA replication licensing in the mouse zygote. Biology of reproduction 10 22674395
2010 Global expression studies in baker's yeast reveal target genes for the improvement of industrially-relevant traits: the cases of CAF16 and ORC2. Microbial cell factories 9 20626860
2018 Identification of a novel trafficking pathway exporting a replication protein, Orc2 to nucleus via classical secretory pathway in Plasmodium falciparum. Biochimica et biophysica acta. Molecular cell research 5 29524523
2014 Dephosphorylation of Orc2 by protein phosphatase 1 promotes the binding of the origin recognition complex to chromatin. Biochemical and biophysical research communications 5 24792176
2025 Anlotinib enhances the efficacy of KRAS-G12C inhibitors through c-Myc/ORC2 axis inhibition in non-small cell lung cancer. Cell death & disease 4 40316534
2017 Reversible regulation of ORC2 SUMOylation by PIAS4 and SENP2. Oncotarget 4 29050267
2024 The ORFIUS complex regulates ORC2 localization at replication origins. NAR cancer 3 38288445
2002 The binding of ORC2 to chromatin from terminally differentiated cells. Experimental cell research 3 11900493
2019 Structural insight into the unique dsDNA binding topology of the human ORC2 wing helix domain. The FEBS journal 2 30963726
2014 Protein phosphatase 1 dephosphorylates Orc2. Biochemical and biophysical research communications 2 24732362
2011 In vivo binding of Orc2 to a region of the chicken lysozyme GAS41 origin containing multiple Sp1-binding sites. DNA and cell biology 2 21879882
2025 Regulation of epigenetics and chromosome structure by human ORC2. bioRxiv : the preprint server for biology 1 39829907
2025 Regulation of epigenetics and chromosome structure by human ORC2. Cell reports 0 40504688
2022 Resonance assignments of the ORC2-WH domain of the human ORC protein. Biomolecular NMR assignments 0 35951249

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