Affinage

Showing IREB2 — IRP2 is a alias.

IREB2

Iron-responsive element-binding protein 2 · UniProt P48200

Length
963 aa
Mass
105.1 kDa
Annotated
2026-06-10
100 papers in source corpus 36 papers cited in narrative 34 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 9/9 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

IREB2 (IRP2) is a cytosolic RNA-binding protein that serves as a central post-transcriptional regulator of cellular and systemic iron homeostasis by binding iron-responsive elements (IREs) in target mRNAs: occupancy of 5'-UTR IREs represses translation (ferritin) while binding of 3'-UTR IREs stabilizes transcripts such as TfR1 (PMID:7890603, PMID:7622457, PMID:15956281). IRP2 binds ferritin IREs with molar efficacy equal to IRP1 but, unlike IRP1, retains RNA-binding activity after alkylation, marking it as a distinct iron sensor (PMID:7890603, PMID:7622457). Its RNA-binding output is governed chiefly by iron- and oxygen-dependent control of its own abundance: under iron- and oxygen-replete conditions IRP2 is targeted for ubiquitin-proteasome degradation, and the FBXL5 E3 ligase complex is the dominant route, with FBXL5 using a [2Fe2S] cluster (oxidized state stabilized by ambient oxygen) to recruit IRP2, sterically dislodge it from IRE RNA, and drive its ubiquitination — accounting for hypoxia-induced IRP2 stabilization (PMID:21907140, PMID:32126207). Degradation requires the C-terminal domain 4, which is necessary and, when transferred, sufficient to confer iron sensitivity on IRP1 (PMID:18226225), and is potentiated by heme binding through the heme regulatory motif (Cys201/His204) that licenses oxidative modification (PMID:16039587); degradation also depends on 2-oxoglutarate-dependent oxygenase activity (PMID:14729944), while raising the labile iron pool is the proximal trigger (PMID:16479012). Additional E3 ligase input from TRIM28, which ubiquitinates IRP2 at K877, and deubiquitinase OTUD3 tune IRP2 levels (PMID:41797698). Beyond protein stability, suppression of iron-sulfur cluster synthesis activates IRP2 RNA binding independently of protein level, FBXL5, and IRP1 (PMID:34039609), and IRP2 transcription is driven by c-MYC and by EpoR/Jak2/Stat5 signaling in erythroid cells (PMID:9924025, PMID:18694996). In vivo, IRP2 is the dominant regulator of erythroid TfR1 and systemic iron distribution, and its loss causes microcytic anemia, motor neuron degeneration with mitochondrial dysfunction, and beta-cell dysfunction through functional iron starvation (PMID:15956281, PMID:22003390, PMID:31941883). Biallelic loss-of-function IREB2 variants in humans cause functional iron deficiency, mitochondrial dysfunction, and neurological disease, with phenotypes reversed by restoring IREB2 expression (PMID:35602653, PMID:39587636, PMID:41234066). IRP2 additionally regulates an expanding set of 3'-UTR IRE-bearing transcripts (Mdm2, TP73, TP53) and HIF2α/EPO, where an IRP2-impairing IRE bulge uridine renders HIF2α regulation IRP1-selective (PMID:31907996, PMID:32304229, PMID:38936518, PMID:39316647).

Mechanistic history

Synthesis pass · year-by-year structured walk · 29 steps
  1. 1995 High

    Established that IRP2 is a bona fide IRE-binding regulator that represses translation via 5'-IREs and stabilizes mRNA via 3'-IREs, distinguishing it biochemically from IRP1.

    Evidence in vitro translation assay and EMSA with purified recombinant IRP2

    PMID:7622457 PMID:7890603

    Open questions at the time
    • In vitro repression did not resolve which endogenous transcripts dominate in vivo
    • Structural basis of IRE recognition not defined
  2. 1995 Medium

    Showed IRP2 is regulated through protein accumulation/degradation rather than the post-translational switch used by IRP1, predicting an independent stability regulator.

    Evidence cycloheximide/actinomycin D treatment with immunoblotting in Ltk- cells

    PMID:7544791

    Open questions at the time
    • The proposed protein controlling IRP2 stability was not identified
    • Pharmacological inhibition is indirect
  3. 1999 Medium

    Connected IRP2 to proliferative signaling by identifying it as a c-MYC transcriptional target that expands the intracellular iron pool.

    Evidence c-MYC gain/loss-of-function and transformation assays

    PMID:9924025

    Open questions at the time
    • Direct promoter binding by c-MYC at IREB2 not shown
    • Single lab
  4. 2003 Medium

    Proposed HOIL-1 as an E3 ligase recognizing heme-oxidized IRP2, offering a mechanism linking heme/iron status to IRP2 turnover.

    Evidence co-IP, ubiquitination and cell-based degradation assays

    PMID:12629548

    Open questions at the time
    • Later contested as non-obligate for iron-mediated degradation
    • Single lab
  5. 2004 High

    Reframed the degradation mechanism by showing it depends on 2-oxoglutarate-dependent oxygenase activity and not the previously proposed cysteine-oxidation domain.

    Evidence deletion/cysteine-mutant analysis with DMOG inhibition in H1299 cells

    PMID:14729944

    Open questions at the time
    • The relevant oxygenase was not identified
    • Direct hydroxylation of IRP2 not demonstrated
  6. 2004 Medium

    Proposed S-nitrosylation at C178 as a route for NO-induced IRP2 degradation.

    Evidence S-nitrosylation assay, C178S mutagenesis and ubiquitination assay in RAW 264.7 cells

    PMID:14673166

    Open questions at the time
    • Later contested by data showing NO-donor degradation is iron-mediated and C178-independent
    • Single lab
  7. 2005 High

    Defined the heme regulatory motif (Cys201/His204) as the heme-sensing element whose oxidative modification triggers degradation and HOIL-1 recognition.

    Evidence spectroscopic heme-binding studies, site-directed mutagenesis, degradation assays

    PMID:16039587

    Open questions at the time
    • Relationship of HRM-dependent degradation to FBXL5 pathway not resolved
    • In vivo relevance of HRM not tested
  8. 2005 High

    Demonstrated in vivo that IRP2 is the dominant stabilizer of erythroid TfR1 mRNA and governs systemic iron distribution.

    Evidence Irp2-/- knockout mice with hematology and TfR1 mRNA quantification

    PMID:15956281

    Open questions at the time
    • Cell-autonomous vs systemic contributions not fully separated
    • Degradation machinery in vivo not addressed
  9. 2005 Medium

    Showed NO can stabilize IRP2 and act as a diffusible intercellular iron-signaling cue, complicating the earlier S-nitrosylation degradation model.

    Evidence NO-donor treatment and co-culture with NOS II-expressing cells, proteasome assays

    PMID:15684386

    Open questions at the time
    • Mechanistic reconciliation of NO stabilizing vs degrading IRP2 unresolved
    • Single lab
  10. 2006 High

    Resolved the NO controversy by showing SNP-induced IRP2 degradation acts by raising the labile iron pool, not via C178 S-nitrosylation.

    Evidence C178S/Delta73 mutants, calcein labile-iron measurement, inhibitor dissection in H1299 cells

    PMID:16479012

    Open questions at the time
    • Did not identify the iron-sensing degradation machinery
    • Single cell-line context
  11. 2007 Medium

    Challenged HOIL-1 as the obligate IRP2 E3 ligase by showing its knockdown does not affect endogenous iron-mediated IRP2 degradation.

    Evidence reciprocal co-IP and HOIL-1 siRNA knockdown in HEK293 cells

    PMID:17822790

    Open questions at the time
    • Did not identify the true iron-dependent E3 ligase
    • Cell-type-specific roles not excluded
  12. 2008 High

    Mapped the C-terminal domain 4 as necessary and transferable for iron-dependent degradation, localizing the degron.

    Evidence truncation and IRP1/IRP2 domain-swap chimeras in H1299 cells

    PMID:18226225

    Open questions at the time
    • The recognizing ligase was still unknown
    • Structural detail of domain 4 not defined
  13. 2008 High

    Linked erythropoietin signaling to iron metabolism by identifying IRP2 as a direct Stat5 transcriptional target whose loss causes microcytic anemia.

    Evidence Stat5-/- mice, ChIP, reporter assays, TfR1 surface flow cytometry

    PMID:18694996

    Open questions at the time
    • Contribution relative to c-MYC-driven transcription unclear
    • Direct vs indirect effects on TfR1 not fully separated
  14. 2008 High

    Provided proof-of-principle rescue showing IRP1 activation by Tempol can compensate for IRP2 loss in brain, restoring iron homeostasis and neuromuscular function.

    Evidence dietary Tempol in Irp2-/- mice, IRP1 IRE-binding and brain TfR1/ferritin assays

    PMID:18685102

    Open questions at the time
    • Long-term and disease-stage efficacy not established
    • Mechanism of Tempol on IRP1 cluster not fully defined
  15. 2010 Medium

    Implicated IRP2 in tumor growth in a manner dependent on its unique 73-aa insert, linking iron handling to proliferation.

    Evidence tetracycline-inducible IRP2 and Delta73 in nude mouse xenografts

    PMID:20405006

    Open questions at the time
    • Mechanistic role of the 73-aa insert in tumorigenesis unresolved
    • Single lab
  16. 2011 High

    Established FBXL5 as the physiologically dominant IRP2 degradation pathway through genetic epistasis, since Irp2 ablation rescues Fbxl5-/- iron-overload lethality.

    Evidence Fbxl5-/- and Fbxl5-/-;Irp2-/- mice with iron-diet challenge

    PMID:21907140

    Open questions at the time
    • Molecular basis of FBXL5 iron/oxygen sensing not yet structural
    • Tissue-specific contributions only partially mapped
  17. 2011 High

    Defined the neurodegenerative consequences of IRP2 loss as functional iron starvation with mitochondrial complex dysfunction, partially rescuable by IRP reactivation or ferritin reduction.

    Evidence Irp2-/- mice, mitochondrial complex assays, genetic ferritin reduction, Tempol

    PMID:22003390

    Open questions at the time
    • Selective neuronal vulnerability not fully explained
    • Link to human disease still pending at this point
  18. 2002 Medium

    Identified YB-1 as a direct iron-dependent IRP2 partner that modulates IRE-dependent translational control.

    Evidence yeast two-hybrid, co-IP, EMSA and in vitro translation reporter

    PMID:12192037

    Open questions at the time
    • In vivo significance of YB-1 modulation unknown
    • Single lab
  19. 2020 High

    Provided the structural and biophysical basis for iron/oxygen sensing, showing FBXL5's oxidized [2Fe2S] cluster organizes IRP2 recruitment and sterically displaces it from IRE RNA.

    Evidence cryo-EM of IRP2-FBXL5-SKP1, EPR spectroscopy, ubiquitination and RNA competition assays

    PMID:32126207

    Open questions at the time
    • Kinetics of cluster oxidation in cells not measured
    • Non-FBXL5 ligase contributions not addressed
  20. 2020 High

    Extended IRP2's physiological reach to beta-cell function by linking iron starvation to defective Fe-S enzyme Cdkal1 and proinsulin misreading.

    Evidence Irp2-/- mice, tRNA modification mass spectrometry, iron-repletion rescue

    PMID:31941883

    Open questions at the time
    • Human diabetes relevance not established
    • Other Fe-S enzymes affected not catalogued
  21. 2021 High

    Showed IRP2 RNA-binding can be activated by ISC synthesis suppression independently of protein level, FBXL5, and IRP1, expanding sensing beyond degradation control.

    Evidence ISC inhibition, IRP1/IRP2 double-deletion cells, direct RNA-binding assays

    PMID:34039609

    Open questions at the time
    • Molecular switch converting ISC status to IRP2 binding undefined
    • In vivo significance at tissue O2 not fully tested
  22. 2020 Medium

    Placed IRP2 within a tumor-suppressor/aging axis by showing it destabilizes TP73 mRNA via a 3'-UTR IRE downstream of FDXR/FDX2.

    Evidence MEFs, FDXR/IRP2 siRNA, 3'UTR IRE reporter assays

    PMID:32304229

    Open questions at the time
    • Physiological setting where this axis dominates unclear
    • Single lab
  23. 2019 Medium

    Identified Mdm2 mRNA as a 3'-UTR IRE target stabilized by IRP2, coupling iron regulation to proliferation control.

    Evidence ectopic IRP2, siRNA/CRISPR KO, IRE-3'UTR stability and reporter assays

    PMID:31907996

    Open questions at the time
    • In vivo relevance of IRP2-Mdm2 axis untested
    • Single lab
  24. 2024 Medium

    Added TP53 as an IRP2-upregulated IRE target and implicated IRP2-driven iron deposition in dopaminergic neuron ferroptosis via SLC7A11-ALOX12.

    Evidence AAV IRP2 overexpression in substantia nigra, MPTP PD model, TP53 3'UTR IRE assay

    PMID:38936518

    Open questions at the time
    • Apparent opposing effects on p73 vs p53 not reconciled
    • Single lab
  25. 2025 High

    Explained divergent IRP1 vs IRP2 erythropoietic phenotypes by an HIF2α IRE bulge uridine that selectively impairs IRP2 binding.

    Evidence IRP1-/-/IRP2-/- mice and in vitro binding with wild-type/mutant IREs, EPO measurement

    PMID:39316647

    Open questions at the time
    • Generalizability of IRE sequence selectivity across targets not mapped
    • Structural basis of bulge discrimination not solved
  26. 2024 High

    Demonstrated human disease causality of biallelic IREB2 loss-of-function through patient cells and lentiviral rescue of iron metabolism defects.

    Evidence patient-derived lymphoblasts, ferritin/TfR1 immunoblot, mitochondrial assays, lentiviral rescue

    PMID:35602653

    Open questions at the time
    • Genotype-phenotype spectrum across patients not fully defined
    • Tissue-specific vulnerability mechanisms incomplete
  27. 2024 High

    Modeled a specific IREB2 missense variant (D826V) causing destabilization, synaptic dysfunction, and behavioral deficits rescuable by proteasome inhibition.

    Evidence CRISPR knock-in mice, behavioral and LTP testing, proteasome-inhibitor rescue, patient cells

    PMID:39587636 PMID:41234066

    Open questions at the time
    • Why D826V destabilizes IRP2 mechanistically not fully defined
    • Translation to patient therapy untested
  28. 2024 Medium

    Identified additional inputs to IRP2 stability and binding via upstream regulators G3BP1 (suppressing FBXL5 translation) and FGFR1 (stabilizing IRP2), tying IRP2 to ferroptosis and cancer iron expansion.

    Evidence co-IP, siRNA, ubiquitination, labile iron and IRE-binding assays in multiple cell lines

    PMID:38118197 PMID:39154074

    Open questions at the time
    • Direct vs indirect effects on IRP2 stability not fully separated
    • Single lab per axis
  29. 2026 High

    Defined TRIM28 as a direct K48-linked E3 ligase ubiquitinating IRP2 at K877, controlling cardiomyocyte iron uptake and ferroptosis.

    Evidence co-IP/MS, ubiquitinome profiling, K877 mutagenesis, cardiomyocyte-specific TRIM28 mouse models

    PMID:41797698

    Open questions at the time
    • Relationship of TRIM28 to FBXL5 pathway hierarchy unresolved
    • Iron/oxygen dependence of TRIM28 targeting not addressed

Open questions

Synthesis pass · forward-looking unresolved questions
  • How the multiple, partly conflicting IRP2 stability inputs (FBXL5, TRIM28, OTUD3, heme/HRM oxidation, ISC sensing, NO) are hierarchically integrated across tissues and oxygen tensions remains unresolved.
  • No unified model ranks the relative contribution of each ligase/DUB in a given tissue
  • Crosstalk between protein-level control and ISC-driven RNA-binding activation undefined
  • Full IRE-target repertoire and sequence determinants of IRP2 vs IRP1 selectivity incomplete

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003723 RNA binding 6 GO:0045182 translation regulator activity 3 GO:0140299 molecular sensor activity 3
Localization
GO:0005829 cytosol 2
Pathway
R-HSA-8953854 Metabolism of RNA 4 R-HSA-382551 Transport of small molecules 3 R-HSA-392499 Metabolism of proteins 3 R-HSA-5357801 Programmed Cell Death 3
Partners
FBXL5G3BP1HOIL-1OTUD3SKP1TRIM28YB-1
Complex memberships
FBXL5-SKP1 (CRL2) E3 ubiquitin ligase complex (as substrate)

Evidence

Reading pass · 34 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1995 IRP2 binds to iron-responsive elements (IREs) in the 5' or 3' UTRs of target mRNAs; binding to 5'-IREs inhibits translation while binding to 3'-IREs stabilizes mRNA. IRP2 was shown to inhibit translation of ferritin mRNAs in vitro with molar efficacy equal to IRP1, and unlike IRP1, is not inactivated for RNA binding by alkylation with N-ethylmaleimide. In vitro translation assay, gel retardation (EMSA), recombinant protein purification The Journal of biological chemistry High 7622457 7890603
1995 IRP2 activation requires de novo protein synthesis (new IRP2 protein accumulation), in contrast to IRP1 which is regulated post-translationally. Iron-induced inactivation/degradation of IRP2 also requires translation of another protein, indicating an independent regulator controls IRP2 stability. Translation inhibitor (cycloheximide) and transcription inhibitor (actinomycin D) treatment, immunoblot analysis in Ltk- cells The Journal of biological chemistry Medium 7544791
1999 c-MYC transcriptionally stimulates IRP2 expression, increasing the intracellular iron pool, as part of a coordinated regulation of iron-controlling genes required for cell proliferation and transformation. Gene expression analysis, c-MYC gain/loss-of-function in cell transformation assays Science (New York, N.Y.) Medium 9924025
1999 Aluminum stabilizes IRP2 by interfering with iron-catalyzed oxidation of IRP2, preventing its degradation and thereby perturbing iron metabolism. IRE binding activity assay, in vitro oxidation assay with aluminum treatment FEBS letters Medium 10580122
2003 The RING finger protein HOIL-1 (heme-oxidized IRP2 ubiquitin ligase-1) functions as an E3 ubiquitin ligase for oxidized IRP2. Heme binds IRP2 in iron-rich cells, generating oxidative modification of IRP2 that serves as a recognition signal for HOIL-1-mediated ubiquitination and degradation. Co-immunoprecipitation, ubiquitination assay, cell-based degradation assay Nature cell biology Medium 12629548
2004 Iron-dependent IRP2 degradation involves 2-oxoglutarate-dependent oxygenase activity, similar to HIF-1alpha degradation. The previously proposed 73-amino-acid degradation domain with cysteine oxidation is not required, as deletion of this domain or triple cysteine mutants remain sensitive to iron-mediated degradation. Dimethyl-oxalylglycine (a 2-oxoglutarate-dependent oxygenase inhibitor) stabilizes IRP2. Mutagenesis of IRP2 expressed in H1299 cells, pharmacological inhibition, proteasome assays Molecular and cellular biology High 14729944
2004 S-nitrosylation of IRP2 at cysteine C178 by nitric oxide leads to IRP2 ubiquitination and proteasomal degradation. Mutation C178S prevents NO-mediated degradation of IRP2. In vitro and in vivo S-nitrosylation assay, site-directed mutagenesis, ubiquitination assay, proteasome inhibitor treatment in RAW 264.7 cells Molecular and cellular biology Medium 14673166
2005 IRP2 degradation is triggered by heme-mediated oxidation involving the heme regulatory motif (HRM). Cys201 binds ferric heme and His204 is a ferrous heme binding site; both residues are critical for IRP2 degradation and recognition by HOIL-1 ubiquitin ligase. The HRM specifically contributes to oxidative modification and iron concentration sensing after iron is integrated into heme. Spectroscopic studies (heme binding), site-directed mutagenesis, cell-based degradation assays Molecular cell High 16039587
2005 IRP2 deficiency in mice causes microcytic anemia and altered body iron distribution due to failure to stabilize TfR1 mRNA in bone marrow erythroid cells, linking IRP2 to systemic iron homeostasis and erythropoiesis. Irp2-/- knockout mouse model, hematological analysis, TfR1 mRNA quantification in bone marrow Blood High 15956281
2005 Nitric oxide (NO) stabilizes IRP2 protein by inhibiting its proteasomal degradation, without compromising overall proteasome activity. NO produced in neighboring cells can stabilize IRP2 by passive diffusion, acting as an intercellular signal. SNAP NO-donor treatment, HA-tagged IRP2 stability assay, co-culture system with NO synthase II-expressing cells, proteasome activity assay in B6 fibroblasts and H1299 cells Molecular and cellular biology Medium 15684386
2006 Sodium nitroprusside (SNP) promotes IRP2 degradation by donating iron into cells and raising the labile iron pool (>4-fold increase), rather than via S-nitrosylation at C178. IRP2 mutants lacking C178 or the 73-aa domain remain sensitive to SNP-induced degradation, and SNP-induced degradation shares the same mechanism as ferric ammonium citrate-induced degradation. IRP2 mutagenesis (C178S, Delta73), calcein-based labile iron pool measurement, pharmacological inhibitors (actinomycin D, cycloheximide, succinylacetone, DMOG) in H1299 cells Molecular and cellular biology High 16479012
2007 HOIL-1 is not required for iron-mediated IRP2 degradation in HEK293 cells. Although HOIL-1 and IRP2 interact via the 73-aa domain, the interaction is not iron-dependent, does not enhance the rate of iron-mediated IRP2 degradation, and siRNA knockdown of HOIL-1 has no effect on endogenous IRP2 degradation. Co-immunoprecipitation, siRNA knockdown of HOIL-1, iron-dependent degradation assay, stable HOIL-1 expression in HEK293 cells Biochimica et biophysica acta Medium 17822790
2008 The C-terminal domain 4 (amino acids 719-963) of IRP2 is necessary but not sufficient for iron-dependent degradation. Deletion mutants lacking this region are stable even in iron-replete cells; transfer of IRP2 domain 4 to IRP1 confers iron sensitivity on the chimeric protein, while substituting IRP1 domain 4 into IRP2 stabilizes the chimeric protein. Systematic mutagenesis (truncation and domain-swap mutants), expression in H1299 cells, iron treatment stability assays BMC molecular biology High 18226225
2008 Stat5 directly transcriptionally regulates IRP2 expression in erythroid cells, linking EpoR/Jak2/Stat5 signaling to iron metabolism. Loss of Stat5 reduces IRP2 transcription, which in turn reduces TfR1 mRNA stability and TfR1 cell surface levels, causing microcytic anemia. Stat5-/- mouse model, chromatin immunoprecipitation (ChIP), transcriptional reporter assays, flow cytometry for TfR1 surface levels Blood High 18694996
2008 Tempol, a stable nitroxide, attenuates neuromuscular impairment in IRP2-/- mice by disassembling the iron-sulfur cluster of IRP1, converting it from cytosolic aconitase to IRE-binding form, which stabilizes TfR1 transcript and represses ferritin synthesis in the brain, partially restoring iron homeostasis. IRP2-/- mouse dietary Tempol supplementation, IRP1 IRE-binding activity assay, TfR1 mRNA and ferritin protein measurements in brain tissues Proceedings of the National Academy of Sciences of the United States of America High 18685102
2010 IRP2 promotes tumor xenograft growth in a manner dependent on its unique 73-amino acid insert; the deletion mutant IRP2-Delta73 fails to promote tumor growth. IRP2 overexpression increases TfR1 levels in xenografts, along with elevated c-MYC and ERK1/2 phosphorylation. Tetracycline-inducible IRP2 expression in nude mouse xenograft model, IRP2-Delta73 deletion mutant, Western blot for TfR1 and signaling proteins, microarray analysis PloS one Medium 20405006
2011 FBXL5-mediated degradation of IRP2 is essential for iron homeostasis in vivo. Fbxl5-/- mice die in utero with excessive iron accumulation, and this lethality is rescued by additional ablation of IRP2, demonstrating that impaired IRP2 degradation is primarily responsible for the phenotype. Fbxl5-/- knockout mice, Fbxl5-/-;Irp2-/- double-knockout rescue, liver-specific Fbxl5 deletion, high-iron diet challenge Cell metabolism High 21907140
2011 IRP2 deficiency causes lower motor neuron degeneration with spinal cord axonopathy and mitochondrial dysfunction (decreased Complex I and II activities) due to functional iron starvation from misregulation of TfR1 and ferritin. Restoring IRP activity (via Tempol activating IRP1) or reducing ferritin expression genetically partially rescues motor neuron degeneration. Irp2-/- mouse model, mitochondrial complex activity assays, histological analysis of spinal cord, genetic ferritin reduction, Tempol dietary rescue PloS one High 22003390
2002 YB-1 (Y-box-binding protein) directly interacts with IRP2 in the presence of high iron concentrations. YB-1 reduces IRP2-mRNA complex formation with ferritin IRE and, when co-administered with IRP2, abrogates the translational inhibition exerted by either protein alone, providing a novel translational control mechanism through IRE. Two-hybrid screen, in vitro binding assay, co-immunoprecipitation, RNA gel shift assay (EMSA), in vitro translation assay with luciferase reporter Molecular and cellular biology Medium 12192037
2020 The C-terminal substrate-binding domain of FBXL5 harbors a [2Fe2S] cluster in the oxidized state that organizes the C-terminal loop responsible for recruiting IRP2. The oxidized state of the cluster maintained by ambient oxygen promotes FBXL5 binding to IRP2, explaining hypoxia-induced IRP2 stabilization. FBXL5 also sterically dislodges IRP2 from IRE RNA to facilitate its ubiquitination and proteasomal turnover. Cryo-EM structure of IRP2-FBXL5-SKP1 complex, EPR spectroscopy of [2Fe2S] cluster, ubiquitination assay, RNA binding competition assay Molecular cell High 32126207
2020 IRP2 loss in mice causes functional iron deficiency in pancreatic beta cells due to dysregulation of TfR1 and ferritin. This impairs Fe-S cluster biosynthesis, reducing the function of Cdkal1 (an Fe-S cluster enzyme), which catalyzes tRNA methylthiolation (ms2t6A37 in tRNALysUUU), leading to proinsulin misreading and impaired insulin processing. Iron repletion normalizes tRNA modification and restores insulin content. Irp2-/- mouse model, tRNA modification mass spectrometry, iron supplementation rescue, metabolic phenotyping Nature communications High 31941883
2021 ISC (iron-sulfur cluster) synthesis suppression can activate IRP2 binding to target mRNAs independent of IRP1, FBXL5, and changes in IRP2 protein level, at tissue-level O2 concentrations. Deletion of both IRP1 and IRP2 abolishes the iron-starvation response, preventing its activation by ISC synthesis inhibition. ISC synthesis inhibition, IRP1/IRP2 double-deletion cell lines, RNA binding activity assay, iron homeostasis measurements Science advances High 34039609
2020 Loss of FDXR increases IRP2 expression (via FDX2), which then binds to an iron response element in the 3'UTR of TP73 mRNA and destabilizes it, repressing p73 protein expression. This FDXR-FDX2-IRP2-TP73 axis regulates aging and tumor suppression. Genetically modified mouse embryonic fibroblasts, siRNA knockdown of FDXR/IRP2, 3'UTR IRE reporter assay, immunoblotting in multiple human cancer cell lines The Journal of pathology Medium 32304229
2019 IRP2 stabilizes Mdm2 mRNA by binding to an iron response element (IRE) in the 3'UTR of Mdm2 mRNA, increasing Mdm2 expression independently of p53. IRP2 knockdown/knockout reduces Mdm2 expression, and IRP2-regulated Mdm2 is required for IRP2-mediated cell proliferation. IRP2 ectopic expression, siRNA/CRISPR knockout in human cancer cells and MEFs, IRE-3'UTR mRNA stability assay, luciferase reporter FASEB journal Medium 31907996
2022 Biallelic loss-of-function IREB2 variants in human patients cause disrupted post-transcriptional regulation of iron metabolism genes (altered ferritin and TfR1 expression), functional iron deficiency, and mitochondrial dysfunction in patient-derived lymphoblasts. Lentiviral restoration of IREB2 expression reverses iron metabolism abnormalities, confirming IRP2 is essential for iron homeostasis in humans. Patient-derived lymphoblast cell line, western blot for ferritin/TfR1, mitochondrial function assay, lentiviral rescue of IREB2 expression Brain communications High 35602653
2023 G3BP1 stabilizes IRP2 by binding to and suppressing translation of FBXL5 mRNA (the E3 ligase component that mediates IRP2 ubiquitination), thereby preventing IRP2 proteasomal degradation, elevating cellular labile iron, and promoting ferroptosis in response to sodium arsenite. Co-immunoprecipitation, siRNA knockdown of G3BP1/FBXL5, IRP2 ubiquitination assay, labile iron measurement, ferroptosis assays in HEK293/MEF/HT1080 cells and mouse model Journal of hazardous materials Medium 38118197
2024 IRP2 post-transcriptionally upregulates TP53 by binding to an IRE in the 3'UTR of TP53 mRNA, increasing p53 protein levels. Elevated IRP2 in substantia nigra increases TFR1 expression, causes iron deposits and dopaminergic neuronal loss, and induces ferroptosis through the SLC7A11-ALOX12 pathway independently of GPX4. AAV-mediated IRP2 overexpression in mouse substantia nigra, MPTP-induced PD model, immunofluorescence, western blot, IRE-binding assay for TP53 3'UTR Free radical biology & medicine Medium 38936518
2025 IRP2 deficiency reduces expression of HIF2α and its transcriptional target EPO, compromising stress erythropoiesis. The HIF2α IRE has a bulge uridine in the upper stem that specifically impairs IRP2 binding but not IRP1 binding, resulting in IRP1-selective regulation of HIF2α and EPO. This explains the distinct erythropoietic phenotypes of IRP1- vs IRP2-deficient mice. IRP1-/- and IRP2-/- mouse models, in vitro IRE binding assays with wild-type and mutant IREs, EPO measurement, erythropoiesis phenotyping Blood High 39316647
2024 The deubiquitinase OTUD3 stabilizes IRP2 protein expression; OTUD3-mediated IRP2 stabilization reduces hippocampal neuron ferroptosis by inhibiting the p53/PTGS2 pathway. Lentiviral OTUD3 overexpression and IRP2 knockdown in cerebral ischemia-reperfusion mouse model and OGD/R neuronal model, co-immunoprecipitation, western blot European journal of medical research Low 39415292
2026 TRIM28 functions as a novel E3 ubiquitin ligase that directly binds IRP2 and promotes K48-linked ubiquitination at the K877 site, leading to IRP2 degradation, reduced TfR1 expression, suppression of intracellular iron uptake, and attenuation of cardiomyocyte ferroptosis during ischemia/reperfusion injury. Co-immunoprecipitation coupled with mass spectrometry, ubiquitinome profiling, AAV9-mediated cardiomyocyte-specific TRIM28 overexpression, cardiomyocyte-specific TRIM28 knockout mice, K877 site mutagenesis Circulation High 41797698
2024 The p.Asp826Val (D826V) missense variant in IRP2 causes significant protein degradation leading to misregulation of intracellular iron homeostasis. In a CRISPR-Cas9 knock-in mouse model, this variant causes reduced IRP2 protein levels, dysregulated iron metabolism, synaptic dysfunction (impaired LTP, increased microglial activation, decreased dendritic spine density), and neurobehavioral deficits. Proteasome inhibitor treatment restores IRP2 expression. CRISPR-Cas9 D826V knock-in mice, behavioral testing (Morris water maze, open field, Y-maze), LTP electrophysiology, proteasome inhibitor rescue, immunoblot in patient-derived cells Orphanet journal of rare diseases / Acta biochimica et biophysica Sinica High 39587636 41234066
2024 FGFR1 stabilizes IRP2 protein through regulation of intracellular protein degradation pathways, leading to increased IRP2 binding to the IRE of TfR1 mRNA, upregulation of TfR1, and expansion of the labile iron pool in prostate cancer cells. Forced IRP2 expression in FGFR1-depleted cells rescues TfR1 expression and tumor growth. FGFR1 knockout (DU145 cells), IRP2 forced expression rescue, IRP2-IRE binding assay (RNA immunoprecipitation), labile iron pool measurement Communications biology Medium 39154074
2018 Leishmania donovani increases IRP2 protein expression and decreases its stability regulator FBXL5 in macrophages/splenocytes, causing increased IRP2 binding to the 5'IRE of ferroportin mRNA and blocking ferroportin translation, thereby retaining iron in macrophages to support intracellular parasite growth. In vitro (J774 macrophages) and in vivo L. donovani infection models, supershift analysis identifying IRP2 involvement, 35S-methionine pulse-chase labeling, FBXL5/ferroportin siRNA knockdown Cellular microbiology Medium 29470856
2024 Diurnal oscillations in IRE-containing transcript regulation in mouse liver are mediated by feeding rhythms rather than the circadian clock. IRP2 protein levels show diurnal variation peaking at the light-dark transition, but ribosome profiling in IRP2-deficient mice reveals that maximum repression of target mRNAs at dark onset still occurs, indicating temporal redundancy with IRP1. IRP2-/- mice, ribosome profiling, liver transcript diurnal time-course analysis, circadian clock mutant mice with controlled feeding Genome biology Medium 38773499

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1999 Coordinated regulation of iron-controlling genes, H-ferritin and IRP2, by c-MYC. Science (New York, N.Y.) 288 9924025
2005 Altered body iron distribution and microcytosis in mice deficient in iron regulatory protein 2 (IRP2). Blood 177 15956281
2021 Iron-sulfur cluster deficiency can be sensed by IRP2 and regulates iron homeostasis and sensitivity to ferroptosis independent of IRP1 and FBXL5. Science advances 158 34039609
2020 FBXL5 Regulates IRP2 Stability in Iron Homeostasis via an Oxygen-Responsive [2Fe2S] Cluster. Molecular cell 145 32126207
2003 Identification of the ubiquitin-protein ligase that recognizes oxidized IRP2. Nature cell biology 143 12629548
2011 The FBXL5-IRP2 axis is integral to control of iron metabolism in vivo. Cell metabolism 142 21907140
2009 Integration of genomic and genetic approaches implicates IREB2 as a COPD susceptibility gene. American journal of human genetics 136 19800047
2023 N6-Methyladenosine-modified circSAV1 triggers ferroptosis in COPD through recruiting YTHDF1 to facilitate the translation of IREB2. Cell death and differentiation 131 36828914
2005 Involvement of heme regulatory motif in heme-mediated ubiquitination and degradation of IRP2. Molecular cell 126 16039587
2013 IRP2 regulates breast tumor growth. Cancer research 118 24285726
1995 Characterization and expression of iron regulatory protein 2 (IRP2). Presence of multiple IRP2 transcripts regulated by intracellular iron levels. The Journal of biological chemistry 117 7622457
2004 Iron-mediated degradation of IRP2, an unexpected pathway involving a 2-oxoglutarate-dependent oxygenase activity. Molecular and cellular biology 106 14729944
1995 Differential modulation of the RNA-binding proteins IRP-1 and IRP-2 in response to iron. IRP-2 inactivation requires translation of another protein. The Journal of biological chemistry 88 7544791
2008 Stat5 regulates cellular iron uptake of erythroid cells via IRP-2 and TfR-1. Blood 86 18694996
2008 Tempol-mediated activation of latent iron regulatory protein activity prevents symptoms of neurodegenerative disease in IRP2 knockout mice. Proceedings of the National Academy of Sciences of the United States of America 82 18685102
1995 Translational repressor activity is equivalent and is quantitatively predicted by in vitro RNA binding for two iron-responsive element-binding proteins, IRP1 and IRP2. The Journal of biological chemistry 79 7890603
2004 S-nitrosylation of IRP2 regulates its stability via the ubiquitin-proteasome pathway. Molecular and cellular biology 73 14673166
2020 Irp2 regulates insulin production through iron-mediated Cdkal1-catalyzed tRNA modification. Nature communications 69 31941883
1992 Molecular cloning, iron-regulation and mutagenesis of the irp2 gene encoding HMWP2, a protein specific for the highly pathogenic Yersinia. Molecular microbiology 65 1552851
1996 Differential regulation of IRP1 and IRP2 by nitric oxide in rat hepatoma cells. Blood 64 8639920
2010 Tumorigenic properties of iron regulatory protein 2 (IRP2) mediated by its specific 73-amino acids insert. PloS one 61 20405006
1998 Converse modulation of IRP1 and IRP2 by immunological stimuli in murine RAW 264.7 macrophages. The Journal of biological chemistry 61 9545264
1993 Chromosomal irp2 gene in Yersinia: distribution, expression, deletion and impact on virulence. Microbial pathogenesis 61 8321119
2011 Iron insufficiency compromises motor neurons and their mitochondrial function in Irp2-null mice. PloS one 57 22003390
2022 MiR-19a suppresses ferroptosis of colorectal cancer cells by targeting IREB2. Bioengineered 53 35599631
2002 Novel translational control through an iron-responsive element by interaction of multifunctional protein YB-1 and IRP2. Molecular and cellular biology 52 12192037
2014 Abnormal brain iron metabolism in Irp2 deficient mice is associated with mild neurological and behavioral impairments. PloS one 48 24896637
2003 The role of endogenous heme synthesis and degradation domain cysteines in cellular iron-dependent degradation of IRP2. Blood cells, molecules & diseases 47 12972033
2004 Targeted mutagenesis of the murine IRP1 and IRP2 genes reveals context-dependent RNA processing differences in vivo. RNA (New York, N.Y.) 44 15208438
2011 The role of IREB2 and transforming growth factor beta-1 genetic variants in COPD: a replication case-control study. BMC medical genetics 42 21320324
2012 Association of IREB2 and CHRNA3 polymorphisms with airflow obstruction in severe alpha-1 antitrypsin deficiency. Respiratory research 38 22356581
1999 Stabilization of iron regulatory protein 2, IRP2, by aluminum. FEBS letters 38 10580122
2002 Increased IRP1 and IRP2 RNA binding activity accompanies a reduction of the labile iron pool in HFE-expressing cells. Journal of cellular physiology 34 11807826
2020 FDXR regulates TP73 tumor suppressor via IRP2 to modulate aging and tumor suppression. The Journal of pathology 33 32304229
2012 Association of IREB2 and CHRNA3/5 polymorphisms with COPD and COPD-related phenotypes in a Chinese Han population. Journal of human genetics 32 22914670
2005 Neurochemical investigations of dopamine neuronal systems in iron-regulatory protein 2 (IRP-2) knockout mice. Brain research. Molecular brain research 32 16051392
1993 Relationship between loss of pigmentation and deletion of the chromosomal iron-regulated irp2 gene in Yersinia pestis: evidence for separate but related events. Infection and immunity 32 8500913
2005 Nitric oxide inhibits the degradation of IRP2. Molecular and cellular biology 31 15684386
2016 The irp2 and fyuA genes in High Pathogenicity Islands are involved in the pathogenesis of infections caused by avian pathogenic Escherichia coli (APEC). Polish journal of veterinary sciences 29 27096784
1995 Virulence-associated fyuA/irp2 gene cluster of Yersinia enterocolitica biotype 1B carries a novel insertion sequence IS1328. FEMS microbiology letters 29 7607411
2015 IREB2 and GALC are associated with pulmonary artery enlargement in chronic obstructive pulmonary disease. American journal of respiratory cell and molecular biology 28 25101718
2014 Presence of fimH, mrkD, and irp2 virulence genes in KPC-2-producing Klebsiella pneumoniae isolates in Recife-PE, Brazil. Current microbiology 27 25085544
2007 Alteration of iron regulatory proteins (IRP1 and IRP2) and ferritin in the brains of scrapie-infected mice. Neuroscience letters 26 17614197
2006 Sodium nitroprusside promotes IRP2 degradation via an increase in intracellular iron and in the absence of S nitrosylation at C178. Molecular and cellular biology 26 16479012
2020 How Oxidation of a Unique Iron-Sulfur Cluster in FBXL5 Regulates IRP2 Levels and Promotes Regulation of Iron Metabolism Proteins. Molecular cell 24 32243827
2019 Zinc induces iron uptake and DMT1 expression in Caco-2 cells via a PI3K/IRP2 dependent mechanism. The Biochemical journal 23 31092704
2024 The involvement of IRP2-induced ferroptosis through the p53-SLC7A11-ALOX12 pathway in Parkinson's disease. Free radical biology & medicine 22 38936518
2013 Association of pro-inflammatory cytokines and iron regulatory protein 2 (IRP2) with Leishmania burden in canine visceral leishmaniasis. PloS one 22 24146743
2023 A regulatory module comprising G3BP1-FBXL5-IRP2 axis determines sodium arsenite-induced ferroptosis. Journal of hazardous materials 20 38118197
2022 Etoposide in combination with erastin synergistically altered iron homeostasis and induced ferroptotic cell death through regulating IREB2/FPN1 expression in estrogen receptor positive-breast cancer cells. Life sciences 20 36442526
2006 Preliminary demonstration of an allelic association of the IREB2 gene with Alzheimer's disease. Journal of Alzheimer's disease : JAD 20 16914832
2005 The pathway for IRP2 degradation involving 2-oxoglutarate-dependent oxygenase(s) does not require the E3 ubiquitin ligase activity of pVHL. Biochimica et biophysica acta 20 15777842
2023 Buddlejasaponin IVb ameliorates ferroptosis of dopaminergic neuron by suppressing IRP2-mediated iron overload in Parkinson's disease. Journal of ethnopharmacology 19 37717841
2022 Disruption of cellular iron homeostasis by IREB2 missense variants causes severe neurodevelopmental delay, dystonia and seizures. Brain communications 19 35602653
2020 Iron overload induced by IRP2 gene knockout aggravates symptoms of Parkinson's disease. Neurochemistry international 18 31904393
2016 Association of IREB2 Gene rs2568494 Polymorphism with Risk of Chronic Obstructive Pulmonary Disease: A Meta-Analysis. Medical science monitor : international medical journal of experimental and clinical research 18 26775557
2008 Iron-dependent degradation of IRP2 requires its C-terminal region and IRP structural integrity. BMC molecular biology 18 18226225
2004 Screening for mutations of the IRP2 gene in Parkinson's disease patients with hyperechogenicity of the substantia nigra. Journal of neural transmission (Vienna, Austria : 1996) 18 15057521
2018 Leishmania donovani inhibits ferroportin translation by modulating FBXL5-IRP2 axis for its growth within host macrophages. Cellular microbiology 17 29470856
2014 Hyperinsulinemia induces hepatic iron overload by increasing liver TFR1 via the PI3K/IRP2 pathway. Journal of molecular endocrinology 17 25385842
2024 Inhibition of IRP2-dependent reprogramming of iron metabolism suppresses tumor growth in colorectal cancer. Cell communication and signaling : CCS 15 39180081
2023 Optimization of Biotinylated RNA or DNA Pull-Down Assays for Detection of Binding Proteins: Examples of IRP1, IRP2, HuR, AUF1, and Nrf2. International journal of molecular sciences 14 36835018
2018 Irp2 Knockout Causes Osteoporosis by Inhibition of Bone Remodeling. Calcified tissue international 14 30191282
2016 Occurrence and analysis of irp2 virulence gene in isolates of Klebsiella pneumoniae and Enterobacter spp. from microbiota and hospital and community-acquired infections. Microbial pathogenesis 14 27133266
2023 Lactobacillus johnsonii L531 Ameliorates Salmonella enterica Serovar Typhimurium Diarrhea by Modulating Iron Homeostasis and Oxidative Stress via the IRP2 Pathway. Nutrients 13 36904126
2019 Oxidative Stress Regulated Iron Regulatory Protein IRP2 Through FBXL5-Mediated Ubiquitination-Proteasome Way in SH-SY5Y Cells. Frontiers in neuroscience 13 30760976
2019 Mdm2 is a target and mediator of IRP2 in cell growth control. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 13 31907996
2018 Genetic variants in FAM13A and IREB2 are associated with the susceptibility to COPD in a Chinese rural population: a case-control study. International journal of chronic obstructive pulmonary disease 13 29872291
2009 On the mechanism of iron sensing by IRP2: new players, new paradigms. Nature chemical biology 13 26190951
2007 HOIL-1 is not required for iron-mediated IRP2 degradation in HEK293 cells. Biochimica et biophysica acta 12 17822790
2019 MiR-935 Promotes Clear Cell Renal Cell Carcinoma Migration and Invasion by Targeting IREB2. Cancer management and research 11 31920398
2014 Iron transport through ferroportin is induced by intracellular ascorbate and involves IRP2 and HIF2α. Nutrients 11 24394537
2003 A high-capacity RNA affinity column for the purification of human IRP1 and IRP2 overexpressed in Pichia pastoris. RNA (New York, N.Y.) 11 12592010
2024 Diurnal control of iron responsive element containing mRNAs through iron regulatory proteins IRP1 and IRP2 is mediated by feeding rhythms. Genome biology 10 38773499
2016 IREB2, CHRNA5, CHRNA3, FAM13A & hedgehog interacting protein genes polymorphisms & risk of chronic obstructive pulmonary disease in Tatar population from Russia. The Indian journal of medical research 10 28474623
2024 Amelioration of nephrotoxicity by targeting ferroptosis: role of NCOA4, IREB2, and SLC7a11 signaling. Brazilian journal of medical and biological research = Revista brasileira de pesquisas medicas e biologicas 9 39383377
2023 Stimulation of Hepatic Ferritinophagy Mitigates Irp2 Depletion-Induced Anemia. Antioxidants (Basel, Switzerland) 9 36978814
2021 Protective Effects of Hif2 Inhibitor PT-2385 on a Neurological Disorder Induced by Deficiency of Irp2. Frontiers in neuroscience 9 34675764
2002 Role of hypolipidemic drug clofibrate in altering iron regulatory proteins IRP1 and IRP2 activities and hepatic iron metabolism in rats fed a low-iron diet. Toxicology and applied pharmacology 9 11969379
2020 Different Associations Between the IREB2 Variants and Chronic Obstructive Pulmonary Disease Susceptibility. Frontiers in genetics 8 33304392
1994 Survey of the irp2 gene among Yersinia pestis strains isolated during several plague outbreaks in northeast Brazil. Memorias do Instituto Oswaldo Cruz 8 7823825
2021 New insights into the construction of wild-type Saba pig-derived Escherichia coli irp2 gene deletion strains. 3 Biotech 7 34466347
2025 A bulge uridine in the HIF2α IRE allows IRP1 but not IRP2 to selectively regulate HIF2α expression and ensuing EPO levels. Blood 6 39316647
2024 FGFR1 governs iron homeostasis via regulating intracellular protein degradation pathways of IRP2 in prostate cancer cells. Communications biology 6 39154074
2024 The deubiquitinase OTUD3 stabilizes IRP2 expression to reduce hippocampal neuron ferroptosis via the p53/PTGS2 pathway to ameliorate cerebral ischemia-reperfusion injury. European journal of medical research 6 39415292
2024 Novel biallelic variants in IREB2 cause an early-onset neurodegenerative disorder in a Chinese pedigree. Orphanet journal of rare diseases 6 39587636
2006 [The role of iron regulatory proteins (IRPs) in the regulation of systemic iron homeostasis: lessons from studies on IRP1 and IRP2 knock out mice]. Postepy higieny i medycyny doswiadczalnej (Online) 5 16819431
2023 β-catenin-IRP2-primed iron availability to mitochondrial metabolism is druggable for active β-catenin-mediated cancer. Journal of translational medicine 4 36703130
2023 Sulforaphane triggers iron overload-mediated ferroptosis in gastric carcinoma cells by activating the PI3K/IRP2/DMT1 pathway. Human & experimental toxicology 4 37201195
2017 Frequency of iss and irp2 genes by PCR method in Escherichia coli isolated from poultry with colibacillosis in comparison with healthy chicken in poultry farms of Zabol, South East of Iran. Polish journal of veterinary sciences 3 28865220
2017 Influence of sodium nitroprusside on expressions of FBXL5 and IRP2 in SH-SY5Y cells. Sheng li xue bao : [Acta physiologica Sinica] 2 28638917
2026 Acupuncture Alleviates Cerebral Ischemia-Reperfusion Injury by Inhibiting Ferroptosis Through the IRP2/IRE Pathway. Clinical and experimental pharmacology & physiology 1 41680076
2025 Nadolol Attenuates Brain Cell Ferroptosis in Ischemic Stroke Rats by Targeting the HOIL-1/IRP2 Pathway. CNS & neurological disorders drug targets 1 39252618
2025 The D826V point mutation in IREB2 causes early-onset neurodegeneration in mice. Acta biochimica et biophysica Sinica 1 41234066
2024 Hsa_circYARS interacts with miR-29a-3p to up-regulate IREB2 and promote laryngeal squamous cell carcinoma progression. Discover oncology 1 39225900
2026 TRIM28 Is an E3 Ligase of IRP2 Suppressing Ischemia/Reperfusion-Induced Myocardial Ferroptosis. Circulation 0 41797698
2025 Targeting iron regulatory protein 2 (IRP2) to disrupt iron metabolism enhances radiosensitivity through mitochondrial dysfunction in breast cancer cells. Cell death discovery 0 40745071
2025 Role of ferroptosis-related IREB2 in the shared genetic etiology between smoking and facial aging: Insights from large-scale genome-wide cross-trait analysis. Computational and structural biotechnology journal 0 40799911
2025 Exploring the Associations Between CHRNA5 and IREB2 Gene Polymorphisms and COPD in the Kazakhstan Population. Biomedicines 0 41007821
2025 Integrative GWAS and Mendelian Randomization Analysis Identifies IREB2 and CD27+ Memory B Cells as Core Drivers of COPD to Lung Cancer Progression. MedComm 0 41377765

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