Affinage

HROB

Homologous recombination OB-fold protein · UniProt Q8N3J3

Length
647 aa
Mass
69.8 kDa
Annotated
2026-06-10
14 papers in source corpus 8 papers cited in narrative 8 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 6/6 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

HROB (C17orf53/MCM8IP) is a nuclear, OB-fold-containing protein that functions as a dedicated loader and activator of the MCM8-MCM9 helicase during homologous recombination–associated DNA synthesis (PMID:31467087, PMID:32528060). It binds ssDNA and directly associates with both the MCM8-9 heterodimer and RPA1, recruiting MCM8-9 to sites of DNA damage to promote HR-associated DNA synthesis (long-tract gene conversion) downstream of RAD51 loading, and these interactions are required for replication fork progression and survival after crosslinking agents (PMID:31467087, PMID:32528060). Biochemically, HROB contacts both MCM8 and MCM9 subunits and binds the heterodimer with highest affinity; it does not affect hexamer assembly but promotes DNA unwinding downstream of loading, likely by coordinating ATP hydrolysis with translocation on branched DNA substrates (PMID:38678026). Cryo-EM analysis indicates HROB drives a rotational rearrangement between the N-terminal DNA-binding and C-terminal AAA+ ATPase domains of MCM8/9 prior to DNA binding, providing a unified model for helicase activation (PMID:42094346). Functionally, HROB is required for interstrand crosslink repair and acts in a pathway epistatic to the Fanconi anemia factor FANCD2, with MCM8/9 operating as a downstream effector of the FA pathway [PMID:32853826, PMID:bio_10.1101_2025.08.07.669127]. Loss of Hrob in mice causes germ cell depletion and infertility consistent with prophase I meiotic arrest, and the HROB-MCM8-9 axis acts redundantly with the HELQ helicase (PMID:31467087).

Mechanistic history

Synthesis pass · year-by-year structured walk · 6 steps
  1. 2019 High

    Established HROB as a recruitment factor that brings the MCM8-9 helicase to DNA damage sites to drive HR-associated DNA synthesis, defining its role downstream of RAD51 and in meiotic recombination.

    Evidence Targeted mouse mutagenesis, genetic epistasis with HELQ, cellular HR and recruitment assays

    PMID:31467087

    Open questions at the time
    • Direct biochemical interaction with MCM8-9 not yet reconstituted at this stage
    • Molecular basis of recruitment to damage sites unresolved
    • How redundancy with HELQ is partitioned mechanistically unclear
  2. 2020 High

    Demonstrated that HROB physically and functionally activates the helicase by binding ssDNA, MCM8-9, and RPA1, with these interactions required for long-tract gene conversion, fork progression, and crosslink survival.

    Evidence In vitro helicase assays with purified proteins, reciprocal Co-IP/pulldown, HR reporters, crosslinker/PARP inhibitor sensitivity in deficient cells

    PMID:32528060 PMID:32853826

    Open questions at the time
    • Structural basis of the HROB–MCM8-9 interface not defined
    • Mechanism by which HROB stimulates unwinding unresolved
    • ICL screen and binding data from a single lab at Medium confidence
  3. 2024 High

    Resolved the HROB–MCM8-9 interaction architecture and showed HROB promotes unwinding downstream of hexamer assembly rather than affecting ring formation, linking activation to ATP hydrolysis at a labile interface.

    Evidence Molecular modeling, ATPase/helicase/binding assays, single-molecule unwinding processivity, interface mutagenesis

    PMID:38678026

    Open questions at the time
    • Atomic-resolution structure of the activated complex not yet captured
    • How HROB couples ATP hydrolysis to translocation only inferred
    • Low processivity in vitro not reconciled with in vivo function
  4. 2024 Low

    Placed HROB expression in S phase co-regulated with replication factors and detected associations with replication machinery, while overexpression suppressed replication.

    Evidence Flow cytometry FRET, cell cycle analysis, ectopic overexpression with replication readout

    PMID:39766854

    Open questions at the time
    • Single FRET method for interaction without orthogonal validation
    • Overexpression phenotype lacks mechanistic resolution
    • Identity of replication-machinery partners not defined
  5. 2025 Medium

    Positioned the HROB-MCM8/9 axis as a downstream effector of the Fanconi anemia pathway, with HROB required for MCM8/9 focus formation at ICLs but dispensable for the FANCD2–MCM8/9 interaction.

    Evidence Co-IP, immunofluorescence focus assays, genetic epistasis (double KO γH2AX/survival)

    PMID:bio_10.1101_2025.08.07.669127

    Open questions at the time
    • Preprint, single lab, not yet peer-reviewed
    • How FANCD2 monoubiquitination licenses MCM9 recruitment mechanistically unresolved
    • Direct vs indirect role of HROB in focus formation not separated
  6. 2026 Medium

    Provided a unified structural model in which HROB drives a rotational rearrangement of MCM8/9 domains prior to DNA binding to enable helicase activation.

    Evidence Cryo-EM with ATP analogs and DNA substrates, conformational state analysis

    PMID:42094346

    Open questions at the time
    • Preprint, single lab, not yet peer-reviewed
    • Functional activation inferred from static conformational states
    • Dynamics of the proposed rotation not directly observed

Open questions

Synthesis pass · forward-looking unresolved questions
  • How HROB integrates helicase activation with RPA exchange, FA-pathway signaling, and replication-coupled regulation in vivo remains to be unified.
  • No high-resolution structure of the fully activated HROB-MCM8/9-DNA complex in a peer-reviewed work
  • Mechanism coupling FANCD2 monoubiquitination to HROB-mediated MCM8/9 loading unresolved
  • Physiological role beyond meiosis and ICL repair not characterized

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003677 DNA binding 2 GO:0060090 molecular adaptor activity 2 GO:0098772 molecular function regulator activity 2
Localization
GO:0005634 nucleus 2
Pathway
R-HSA-73894 DNA Repair 3 R-HSA-69306 DNA Replication 2

Evidence

Reading pass · 8 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2019 HROB (C17orf53) is an OB-fold-containing protein that recruits the MCM8-MCM9 helicase to sites of DNA damage to promote HR-associated DNA synthesis downstream of RAD51. Mice with targeted Hrob mutations are infertile due to germ cell depletion consistent with prophase I meiotic arrest. The HROB-MCM8-MCM9 pathway acts redundantly with the HELQ helicase; cells lacking both HROB and HELQ have severely impaired HR. Targeted mouse mutagenesis, genetic epistasis (HROB/HELQ double KO), cellular HR assays, focus formation/recruitment assays Genes & development High 31467087
2020 HROB (MCM8IP/C17orf53) binds ssDNA and directly associates with both the MCM8-9 helicase complex and RPA1. HROB stimulates the helicase activity of MCM8-9 in vitro, and its interactions with MCM8-9 and RPA are required for HR (specifically long-tract gene conversion downstream of RAD51 loading), replication fork progression, and cellular viability after crosslinking agent treatment. In vitro helicase activity assay with purified proteins, Co-IP/pulldown (HROB–MCM8-9 and HROB–RPA1), HR reporter assays, crosslinker/PARP inhibitor sensitivity assays in MCM8IP-deficient cells Nature communications High 32528060
2020 C17orf53/HROB is a ssDNA- and RPA-binding protein required for interstrand crosslink (ICL) repair and efficient DNA replication; its loss leads to hypersensitivity to ATR inhibition and pronounced ICL repair defects. HROB works together with MCM8/9 to promote cell survival after ICL lesions. Genome-wide ATR-inhibitor fitness screen, multi-omics interaction studies, ssDNA/RPA-binding assays, cellular ICL sensitivity assays, DNA replication rate measurement in C17orf53 KO cells DNA repair Medium 32853826
2024 Biochemical and structural analysis defined the HROB–MCM8-9 interaction interface: HROB makes transient contacts with both MCM8 and MCM9 subunits and binds the MCM8-9 heterodimer with highest affinity. MCM8-9-HROB preferentially binds and unwinds branched DNA structures with low processivity. MCM8-9 assembles into a hexamer from dimers on DNA in the presence of ATP via two alternating interfaces (one stable/obligate heterodimer, one labile mediating hexamer assembly). HROB does not affect ring/hexamer formation but promotes DNA unwinding downstream of loading, likely by coordinating ATP hydrolysis with translocation. The ATPase site at the labile interface contributes disproportionately more to unwinding. Molecular modeling, biochemistry (ATPase assays, helicase assays, binding assays), single-molecule experiments (unwinding processivity), mutational analysis of interfaces Nature communications High 38678026
2024 HROB is a nuclear protein whose level peaks in S phase and is downregulated in quiescence, co-regulated with DNA replication factors. Flow cytometry FRET assays detected associations between HROB and proteins of the DNA replication machinery. Ectopic overexpression of HROB caused an almost complete shutdown of DNA replication. Flow cytometry FRET (protein–protein association), cell cycle analysis, ectopic overexpression with DNA replication readout, structural prediction/homology analysis Genes Low 39766854
2026 Cryo-EM structures of MCM8/9 with DNA, HROB, and ATP analogs revealed that DNA binding induces a rotational rearrangement between N-terminal DNA-binding and C-terminal AAA+ ATPase domains, reorganizing DNA-binding loops into a staircase configuration. HROB associates with both halves of the heterohexamer and drives a similar rotation prior to DNA binding, localizing MCM8/9 to crosslink damage sites and enabling unwinding, providing a unified mechanistic model for MCM8/9 helicase activation by HROB. Cryo-electron microscopy (cryo-EM) with ATP analogs and DNA substrates, structural analysis of conformational states bioRxivpreprint Medium 42094346
2025 HROB is required for MCM8/9 focus formation at ICL damage sites (functional activation context), but HROB is not required for FANCD2 binding to MCM8/9. MCM8/9 physically interacts with FANCD2 through its core domain independently of DNA, and FANCD2 acts upstream of MCM9 recruitment to ICL-induced nuclear foci downstream of FANCD2 monoubiquitination. MCM8/9 and FANCD2 are epistatic (combined loss does not cause additive DNA damage), placing MCM8/9 as a downstream effector of the FA pathway. Co-immunoprecipitation, immunofluorescence focus assays, genetic epistasis (double KO γH2AX and survival assays), MCM8/9 KO cell lines bioRxivpreprint Medium bio_10.1101_2025.08.07.669127
2025 HROB suppresses lung adenocarcinoma cell proliferation by interacting with ZC3HC1 and reducing its phosphorylation at Ser354, which facilitates K27-linked ubiquitination of CCNB1 and its proteasomal degradation, impairing the G2-to-M phase transition. Co-immunoprecipitation (HROB–ZC3HC1 interaction), phosphorylation analysis, ubiquitination assay, proteasomal degradation assay, cell proliferation/tumor growth assays with KO/overexpression Cancer science Low 40654113

Source papers

Stage 0 corpus · 14 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2019 Control of homologous recombination by the HROB-MCM8-MCM9 pathway. Genes & development 66 31467087
2020 MCM8IP activates the MCM8-9 helicase to promote DNA synthesis and homologous recombination upon DNA damage. Nature communications 47 32528060
2021 Meiotic genes in premature ovarian insufficiency: variants in HROB and REC8 as likely genetic causes. European journal of human genetics : EJHG 39 34707299
2020 C17orf53 is identified as a novel gene involved in inter-strand crosslink repair. DNA repair 24 32853826
2024 Mechanism of DNA unwinding by MCM8-9 in complex with HROB. Nature communications 13 38678026
2019 Ways to unwind with HROB, a new player in homologous recombination. Genes & development 11 31575675
2022 Pan-Cancer Analysis Identified Homologous Recombination Factor With OB-Fold (HROB) as a Potential Biomarker for Various Tumor Types. Frontiers in genetics 4 35903352
2025 HROB Induces Lung Adenocarcinoma Progression via ZC3HC1-CCNB1 Axis Regulation and Cell Cycle Dysregulation. Cancer science 1 40654113
2024 HROB Is Implicated in DNA Replication. Genes 1 39766854
2023 Mechanism of DNA unwinding by hexameric MCM8-9 in complex with HROB. Research square 1 37461676
2023 Genetic analysis of novel pathogenic gene HROB in a family with primary ovarian insufficiency. Zhejiang da xue xue bao. Yi xue ban = Journal of Zhejiang University. Medical sciences 1 38105698
2026 Biallelic Germline Inactivation of HROB Causes Primary Gonadal Insufficiency and is Potentially Associated with Colonic Polyposis Predisposition. American journal of medical genetics. Part A 0 41878782
2026 Structural Activation of DNA Unwinding by MCM8/9/HROB. bioRxiv : the preprint server for biology 0 42094346
2023 Mechanism of DNA unwinding by hexameric MCM8-9 in complex with HROB. bioRxiv : the preprint server for biology 0 37398313

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