Affinage

RNF212

Probable E3 SUMO-protein ligase RNF212 · UniProt Q495C1

Length
297 aa
Mass
33.4 kDa
Annotated
2026-06-10
36 papers in source corpus 14 papers cited in narrative 14 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

RNF212 is a dosage-sensitive, limiting regulator of meiotic crossover designation that couples chromosome synapsis to the assembly of crossover-specific recombination complexes (PMID:23396135). It selectively localizes to a subset of recombination sites in a synapsis-dependent manner and stabilizes pro-crossover factors, including the MutSγ complex (MSH4-MSH5), marking and sustaining the pre-crossover state (PMID:23396135, PMID:24891606). Mechanistically, RNF212 acts as a SUMO E3 ligase on meiotic chromosome axes, where RNF212-dependent SUMO conjugation establishes a checkpoint-like state that stalls recombination and renders turnover of a subset of recombination factors dependent on HEI10-mediated ubiquitylation; this SUMO–ubiquitin relay recruits proteasomes to chromosome axes to resolve the limited final set of crossovers (PMID:28059716). The transition from pre-crossover to mature crossover requires CNTD1, which regulates the association between HEI10 and RNF212 (PMID:24891606). RNF212 operates within an interdependent RING-domain E3 ligase network with its paralog RNF212B, which it physically binds and whose chromosomal loading it controls, and both are recruited to recombination sites by HEIP1 acting through HEI10 (PMID:38865271, PMID:41118211). Beyond crossover control, RNF212 mediates oocyte quality control by impeding DSB repair and enabling HORMAD1 association with desynapsing chromosomes, sensitizing oocytes to DSB-induced apoptosis (PMID:30270110). Homozygous loss-of-function variants in human RNF212 cause male meiotic arrest, establishing it as required for human male meiosis (PMID:31125047).

Mechanistic history

Synthesis pass · year-by-year structured walk · 7 steps
  1. 2013 High

    Established that RNF212 is the limiting, dosage-sensitive factor coupling synapsis to crossover designation, answering how synapsis is translated into a defined number of crossovers.

    Evidence Mouse knockout and haploinsufficiency analysis with immunofluorescence of recombination factors on meiotic spreads

    PMID:23396135

    Open questions at the time
    • Molecular enzymatic activity of RNF212 not yet defined
    • Direct biochemical interaction with MutSγ not shown
  2. 2014 High

    Positioned RNF212 within the pathway by showing its prolonged association marks the pre-crossover state and that CNTD1 governs the RNF212–HEI10 transition to mature crossovers.

    Evidence Mouse Cntd1 knockout with quantitative immunofluorescence of RNF212, MutSγ, HEI10, MutLγ foci

    PMID:24891606

    Open questions at the time
    • Does not establish whether CNTD1 acts directly on RNF212
    • Biochemical nature of RNF212 turnover not resolved
  3. 2017 High

    Defined the molecular activity of RNF212 as a SUMO E3 ligase that imposes a checkpoint-like recombination stall coupled to a SUMO–ubiquitin–proteasome relay on chromosome axes.

    Evidence Mouse Rnf212 and Hei10 mutant genetics, SUMO/ubiquitin conjugation assays, and proteasome recruitment on meiotic spreads

    PMID:28059716

    Open questions at the time
    • Specific SUMO substrates of RNF212 not enumerated
    • Structural basis of SUMO ligase activity not determined
  4. 2018 High

    Revealed a second, distinct RNF212 function in oocyte quality control by impeding DSB repair and enabling HORMAD1 loading on desynapsing chromosomes.

    Evidence Mouse Rnf212 knockout with oocyte survival assays, HORMAD1 localization, and DNA damage marker analysis

    PMID:30270110

    Open questions at the time
    • Mechanism linking RNF212 SUMO activity to apoptotic sensitization unclear
    • Whether the same enzymatic activity drives both crossover and quality-control roles not resolved
  5. 2019 Medium

    Extended RNF212 function to humans, showing homozygous loss-of-function causes male meiotic arrest.

    Evidence Gene-panel sequencing with immunohistochemistry/cytochemistry on testicular biopsies from variant carriers

    PMID:31125047

    Open questions at the time
    • Single-lab human cohort
    • Female meiotic consequences in humans not assessed
  6. 2024 High

    Identified RNF212B as a physically interacting paralog whose chromosomal loading depends on RNF212, defining an interdependent two-paralog E3 ligase module.

    Evidence Mouse single/double knockouts, co-IP/pull-down, SUMOylome MS, ubiquitin pull-down, and cytology

    PMID:38865271

    Open questions at the time
    • Direct substrates distinguishing RNF212 vs RNF212B activity not defined
    • Structural basis of the RNF212–RNF212B interaction unknown
  7. 2025 High

    Resolved the division of labor among the crossover-regulating RING proteins RNF212, RNF212B, and HEI10, and identified HEIP1 as the upstream factor recruiting RNF212/RNF212B via HEI10.

    Evidence Mouse single/double mutants, live imaging, and Heip1 KO with co-IP and immunofluorescence across sexes and stages

    PMID:39761402 PMID:41118211

    Open questions at the time
    • Biochemical hierarchy of recruitment among HEIP1, HEI10, and RNF212 not fully reconstituted
    • Sex-specific differences in localization dynamics mechanistically unexplained

Open questions

Synthesis pass · forward-looking unresolved questions
  • How RNF212's SUMO E3 activity is biochemically reconstituted, which substrates it modifies in vivo, and how the same enzyme partitions between crossover designation and oocyte quality control remain open.
  • No in vitro reconstitution of RNF212 SUMO ligase activity with defined substrates
  • No structural model of RNF212 or its paralog complex
  • Mechanism partitioning crossover versus quality-control functions undefined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0098772 molecular function regulator activity 2 GO:0016740 transferase activity 1 GO:0140096 catalytic activity, acting on a protein 1
Localization
GO:0005694 chromosome 3 GO:0005634 nucleus 1
Pathway
R-HSA-1474165 Reproduction 2 R-HSA-1640170 Cell Cycle 2 R-HSA-392499 Metabolism of proteins 1

Evidence

Reading pass · 14 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2013 Mouse RNF212 is essential for meiotic crossing-over; it functions to couple chromosome synapsis to the formation of crossover-specific recombination complexes. RNF212 selectively localizes to a subset of recombination sites (an early step in crossover designation) and acts to stabilize meiosis-specific recombination factors including the MutSγ complex (MSH4-MSH5). Haploinsufficiency demonstrates that RNF212 is a dosage-sensitive, limiting factor for crossover control. Mouse knockout and heterozygous loss-of-function combined with immunofluorescence localization of recombination factors on meiotic chromosome spreads Nature genetics High 23396135
2017 RNF212 acts as a SUMO E3 ligase on meiotic chromosome axes; RNF212-dependent SUMO conjugation establishes a checkpoint-like state that stalls recombination by rendering turnover of a subset of recombination factors dependent on HEI10-mediated ubiquitylation. Interdependent localization of SUMO, ubiquitin, and proteasomes along chromosome axes is mediated largely by RNF212 and HEI10. This SUMO–ubiquitin relay recruits proteasomes to chromosome axes to regulate meiotic recombination. Mouse genetics (Rnf212 KO and Hei10 mutants), immunofluorescence, SUMO/ubiquitin conjugation assays on meiotic spreads, proteasome recruitment to chromosomes Science (New York, N.Y.) High 28059716
2014 RNF212 marks pre-crossover intermediates and its prolonged association with recombination sites (along with MutSγ) is a defining feature of the pre-CO state; CNTD1 is required for the transition from pre-CO to mature CO by regulating the association between HEI10 and RNF212 and CO machinery components. Loss of CNTD1 leads to failure to localize MutLγ and HEI10 at designated CO sites and to sustained high levels of MutSγ/RNF212 foci. Mouse Cntd1 knockout, immunofluorescence quantification of RNF212, MutSγ, HEI10, MutLγ foci on meiotic chromosome spreads The Journal of cell biology High 24891606
2018 RNF212 also mediates oocyte quality control: it sensitizes oocytes to DSB-induced apoptosis within a narrow window as chromosomes desynapse. RNF212 is required for HORMAD1 (a negative regulator of inter-sister recombination) to associate with desynapsing chromosomes, and RNF212 impedes DSB repair, retaining a 'memory' of meiotic defects for quality control. Mouse Rnf212 knockout, oocyte survival assays, immunofluorescence for HORMAD1 on desynapsing chromosomes, DNA damage marker analysis Molecular cell High 30270110
2025 RNF212, HEI10, and RNF212B are three distinct mammalian crossover-regulating (COR) RING-domain proteins with divergent spatiotemporal localization dynamics along synapsed chromosomes (including profound differences between spermatocytes and oocytes). Contrasting mutant phenotypes and genetic requirements indicate they play distinct but interdependent functions in regulating meiotic recombination, integrating signals from DNA breaks, homolog synapsis, cell cycle, and incipient crossover sites. Mouse genetics (single and double mutants), live imaging and immunofluorescence tracking of RNF212, HEI10, RNF212B localization dynamics in spermatocytes and oocytes Proceedings of the National Academy of Sciences of the United States of America High 39761402
2024 RNF212B colocalizes and physically interacts with RNF212, forming foci along chromosomes from zygonema onward in a synapsis-dependent, DSB-independent manner. RNF212B foci formation depends genetically on Rnf212 but not on Msh4, Hei10, or Cntd1; unloading of RNF212B at end of pachynema depends on Hei10 and Cntd1. SUMOylome analysis and pull-down assays indicate RNF212B has ubiquitin E3 ligase activity. Double mutants for Rnf212b and Rnf212 are phenotypically identical to Rnf212b single mutants, while double heterozygotes show dosage-dependent CO reduction, indicating functional interplay between paralogs. Mouse knockouts (single and double), co-immunoprecipitation/pull-down, SUMOylome mass spectrometry, immunofluorescence on meiotic spreads, genetic epistasis Proceedings of the National Academy of Sciences of the United States of America High 38865271
2025 HEIP1 directly interacts with HEI10 and orchestrates the recruitment of pro-CO E3 ligases RNF212 and RNF212B (as well as MutSγ complex) to meiotic recombination sites. Loss of HEIP1 impairs RNF212 and RNF212B localization at crossover sites, and recruitment of MutLγ resolution complex. Mouse Heip1 knockout, co-immunoprecipitation (HEIP1–HEI10 interaction), immunofluorescence for RNF212, RNF212B, HEI10, MutLγ on meiotic spreads Proceedings of the National Academy of Sciences of the United States of America High 41118211
2019 Homozygous loss-of-function variants in human RNF212 cause meiotic arrest in males, establishing RNF212 as required for human male meiosis. Meiotic studies of testicular biopsies from mutation carriers confirmed functional consequences consistent with roles defined in mouse knockout models. Next-generation sequencing (gene panel), immunohistochemistry and immunocytochemistry on human testicular biopsies from homozygous RNF212 variant carriers Human reproduction (Oxford, England) Medium 31125047
2014 An ENU-induced nonsense mutation in mouse Rnf212 (repro57 allele) causes male infertility with spermatocyte degeneration at late meiotic prophase, greatly reduced MLH1 (crossover) foci, absent chiasmata, and premature XY dissociation, confirming that RNF212 is required for crossover formation and late prophase progression. ENU mutagenesis screen, histology, MLH1 immunofluorescence on meiotic spreads, genetic mapping of repro57 mutation to Rnf212 Reproduction (Cambridge, England) Medium 25342176
2024 Female repro57 (Rnf212 nonsense mutation) homozygous mutant oocytes exhibit complete aneuploidy with increased kinetochore distances, and derived embryos show significantly reduced morula/blastocyst rates with frequent cytokinesis failure and vacuole formation, indicating that RNF212 loss leads to premature sister chromatid separation that propagates through embryo development. In vitro fertilization of repro57 homozygous mutant oocytes, chromosome spreading and FISH for aneuploidy, immunofluorescence for kinetochore markers, embryo developmental tracking Reproduction (Cambridge, England) Medium 39042717
2016 In C. elegans, the MAP kinase pathway (via MPK-1) coordinates crossover designation with SC protein disassembly; inactivation of MPK-1 at late pachytene is required for SC disassembly from long arms and depends on ZHP-3 (the C. elegans ortholog of RNF212/Zip3) and COSA-1 (CNTD1 ortholog), placing ZHP-3/RNF212 as a required factor downstream of crossover designation for coordinating SC remodeling. C. elegans genetics (ZHP-3/COSA-1 mutants), phosphorylation analysis of SYP-2, MAP kinase pathway manipulation by constitutively active and dominant-negative alleles eLife Medium 26920220
2017 In budding yeast, Zip3 (the RNF212/ZHP-3 ortholog) recruits proteolytic core and regulatory proteasome particles to meiotic chromosomes (together with SC protein Zip1), and proteasome function is required for the coordinated transition entailing SC assembly and stable strand exchange of crossover-designated DSBs, establishing a conserved chromosome-axis-based proteasome recruitment role for this protein family. Yeast genetics, proteasome localization by immunofluorescence, Zip3 mutant analysis for chromosome pairing and crossover formation Science (New York, N.Y.) Medium 28059715
2014 In yeast, Zip3 (RNF212 ortholog) promotes biased cutting of double Holliday junction intermediates toward crossover outcomes, as inferred from genome-wide recombination intermediate signatures in zip3 mutants compared to other ZMM mutants. Genome-wide sequencing of recombination products in yeast mutants (zip3, sgs1, mms4-md) to analyze conversion tracts and crossover/non-crossover signatures PLoS genetics Medium 25329811
2018 In C. elegans, ZHP-4 (an RNF212-like paralog of ZHP-3) cooperates with ZHP-3 to enforce crossover formation at two distinct steps: formation of early joint molecules and transition of late CO intermediates into chiasmata. RING domain mutants show that ZHP-4 is required for localization of both ZHP-3 and ZHP-4 to the synaptonemal complex, and for stabilizing pro-CO factors (MSH-5, RMH-1, COSA-1) at designated CO sites. C. elegans genetics (zhp-4 null and hypomorphic mutants, RING domain mutants), immunofluorescence for pro-CO factors on meiotic chromosomes PLoS genetics Medium 30379819

Source papers

Stage 0 corpus · 36 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2013 RNF212 is a dosage-sensitive regulator of crossing-over during mammalian meiosis. Nature genetics 231 23396135
2008 Sequence variants in the RNF212 gene associate with genome-wide recombination rate. Science (New York, N.Y.) 161 18239089
2017 A SUMO-ubiquitin relay recruits proteasomes to chromosome axes to regulate meiotic recombination. Science (New York, N.Y.) 151 28059716
2016 Conserved Genetic Architecture Underlying Individual Recombination Rate Variation in a Wild Population of Soay Sheep (Ovis aries). Genetics 126 27029733
2009 Genetic analysis of variation in human meiotic recombination. PLoS genetics 126 19763160
2017 Control of meiotic pairing and recombination by chromosomally tethered 26S proteasome. Science (New York, N.Y.) 74 28059715
2016 Comparative analysis of testis transcriptomes associated with male infertility in cattleyak. Theriogenology 68 27865410
2019 Sequencing of a 'mouse azoospermia' gene panel in azoospermic men: identification of RNF212 and STAG3 mutations as novel genetic causes of meiotic arrest. Human reproduction (Oxford, England) 64 31125047
2014 Mammalian CNTD1 is critical for meiotic crossover maturation and deselection of excess precrossover sites. The Journal of cell biology 64 24891606
2018 Impeding DNA Break Repair Enables Oocyte Quality Control. Molecular cell 52 30270110
2014 Controlling meiotic recombinational repair - specifying the roles of ZMMs, Sgs1 and Mus81/Mms4 in crossover formation. PLoS genetics 48 25329811
2018 Identification of a multidimensional transcriptome signature for survival prediction of postoperative glioblastoma multiforme patients. Journal of translational medicine 46 30572911
2016 The MAP kinase pathway coordinates crossover designation with disassembly of synaptonemal complex proteins during meiosis. eLife 38 26920220
2018 A Genomic Region Containing REC8 and RNF212B Is Associated with Individual Recombination Rate Variation in a Wild Population of Red Deer (Cervus elaphus). G3 (Bethesda, Md.) 37 29764960
2022 Integrative RNA profiling of TBEV-infected neurons and astrocytes reveals potential pathogenic effectors. Computational and structural biotechnology journal 31 35685361
2018 Genomic characterization of cervical cancer based on human papillomavirus status. Gynecologic oncology 30 30581036
2018 C. elegans ZHP-4 is required at multiple distinct steps in the formation of crossovers and their transition to segregation competent chiasmata. PLoS genetics 26 30379819
2024 RNF212B E3 ligase is essential for crossover designation and maturation during male and female meiosis in the mouse. Proceedings of the National Academy of Sciences of the United States of America 18 38865271
2017 A Zip3-like protein plays a role in crossover formation in the SC-less meiosis of the protist Tetrahymena. Molecular biology of the cell 15 28100637
2014 An ENU-induced mutation in the mouse Rnf212 gene is associated with male meiotic failure and infertility. Reproduction (Cambridge, England) 15 25342176
2022 Recombination rates in pigs differ between breeds, sexes and individuals, and are associated with the RNF212, SYCP2, PRDM7, MEI1 and MSH4 loci. Genetics, selection, evolution : GSE 14 35596132
2025 Distinct and interdependent functions of three RING proteins regulate recombination during mammalian meiosis. Proceedings of the National Academy of Sciences of the United States of America 12 39761402
2016 Multiple roles of FOXJ3 in spermatogenesis: A lesson from Foxj3 conditional knockout mouse models. Molecular reproduction and development 10 27739607
2023 A pathogenic variant in the uncharacterized RNF212B gene results in severe aneuploidy male infertility and repeated IVF failure. HGG advances 8 37124137
2018 Identification of Serial DNA Methylation Changes in the Blood Samples of Patients with Lung Cancer. Tuberculosis and respiratory diseases 7 30302959
2016 Genome-Wide Association Study of Meiotic Recombination Phenotypes. G3 (Bethesda, Md.) 7 27733454
2017 Association of the common SNPs in RNF212, STAG3 and RFX2 gene with male infertility with azoospermia in Chinese population. European journal of obstetrics, gynecology, and reproductive biology 5 29277047
2024 Chromosomal missegregation and aberrant embryo development in repro57 female mice with Rnf212 homozygous mutation. Reproduction (Cambridge, England) 2 39042717
2025 Common variation in meiosis genes shapes human recombination phenotypes and aneuploidy risk. medRxiv : the preprint server for health sciences 1 40321295
2025 Men with genetic predisposition face greater fertility challenges when exposed to electromagnetic radiation. Molecular biology reports 1 40742570
2026 Common variation in meiosis genes shapes human recombination and aneuploidy. Nature 0 41565805
2025 Exploring the therapeutic effect of melatonin targeting common biomarkers in testicular germ cell tumor, prostate adenocarcinoma, and male infertility: an integrated biology approach. Mammalian genome : official journal of the International Mammalian Genome Society 0 40056207
2025 A variant in RNF212B may contribute to female infertility and recurrent pregnancy loss. HGG advances 0 40259604
2025 HEIP1 orchestrates pro-crossover protein activity during mammalian meiosis. bioRxiv : the preprint server for biology 0 40909735
2025 HEIP1 orchestrates pro-crossover protein activity during mammalian meiosis. Proceedings of the National Academy of Sciences of the United States of America 0 41118211
2018 The Association of rs1670533 Polymorphism in RNF212 Gene With the Risk of Down Syndrome in Young Women. Journal of family & reproductive health 0 30647754

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