Affinage

XIRP2

Xin actin-binding repeat-containing protein 2 · UniProt A4UGR9

Length
3374 aa
Mass
382.3 kDa
Annotated
2026-06-11
61 papers in source corpus 12 papers cited in narrative 12 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 5/5 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

XIRP2 (Xin actin-binding repeat-containing protein 2) is a striated muscle- and hair cell-expressed cytoskeletal protein whose tandem Xin repeats constitute a direct actin-binding motif that crosslinks F-actin into stabilized networks (PMID:15454575). In the heart, XIRP2 is a direct transcriptional target of MEF2A induced by angiotensin II signaling, and its loss attenuates hypertrophic remodeling, fibrosis, and apoptosis following Ang II challenge, placing it in the cardiac stress-response pathway (PMID:17046827, PMID:20093629). At the sarcomere it localizes to the Z-disc/costamere, interacts with alpha-actinin-2, and stabilizes filamin C in premyofibrils (PMID:17046827, PMID:27206985). XIRP2 is required for postnatal maturation of cardiac intercalated discs, where it drives redistribution of junctional components to cardiomyocyte termini and associates with the Nav1.5 and Kv1.5 ion channel subunits to support normal conduction (PMID:23261932, PMID:29306897); compound heterozygous truncating XIRP2 mutations are linked to dilated cardiomyopathy through disrupted intercalated disc integrity (PMID:26656454). In inner ear hair cells, isoform-specific XIRP2 (including a short Xin-repeat-deficient form, XEPLIN) localizes to stereocilia and the cuticular plate and is required for long-term maintenance of stereocilia F-actin architecture and hearing (PMID:25653358, PMID:25772365); a C-terminal mechanosensor domain mediates force-dependent recruitment of XIRP2 to damaged actin sites, where it facilitates monomeric actin enrichment to repair noise-induced stereocilia damage (PMID:37294664).

Mechanistic history

Synthesis pass · year-by-year structured walk · 11 steps
  1. 2004 High

    Established that the defining Xin repeats are themselves a functional actin-binding module, providing the biochemical basis for all later cytoskeletal roles.

    Evidence In vitro co-sedimentation with skeletal muscle actin and repeat-construct transfection in cultured cells

    PMID:15454575

    Open questions at the time
    • Affinity and stoichiometry of binding not quantified
    • Does not address regulation of binding in vivo
    • Structural basis of the actin-bundling activity unresolved
  2. 2006 Medium

    Placed XIRP2 in a defined transcriptional and structural context by identifying it as a MEF2A target localizing to the Z-disc and interacting with alpha-actinin-2.

    Evidence MEF2A knockout mouse microarray, co-IP/pulldown, and muscle immunolocalization

    PMID:17046827

    Open questions at the time
    • alpha-actinin-2 interaction from single lab, no reciprocal validation
    • Functional consequence of the Z-disc interaction untested
    • Direct vs indirect MEF2A regulation not distinguished at the time
  3. 2010 High

    Connected XIRP2 to cardiac stress signaling, showing Ang II induces it via MEF2A and that its reduction alters hypertrophic remodeling.

    Evidence Xirp2 promoter and MEF2A activity assays plus Xirp2 hypomorphic mice under chronic Ang II infusion

    PMID:20093629

    Open questions at the time
    • Mechanism linking actin-binding to remodeling output unclear
    • Hypomorph rather than full null
    • Cell-autonomous vs systemic contributions not separated
  4. 2012 High

    Defined an essential developmental role: XIRP2 drives postnatal redistribution of junctional components to form mature intercalated discs.

    Evidence mXinβ knockout mouse with Western blot, immunofluorescence, fractionation, and epistatic comparison to mXinα-null

    PMID:23261932

    Open questions at the time
    • Molecular trigger of XIRP2 upregulation during redistribution unknown
    • Direct binding partners at the ICD not defined here
    • Mechanism of component redistribution not resolved
  5. 2015 High

    Extended XIRP2 function to a new tissue, demonstrating isoform-specific stereocilia localization and a requirement for long-term hair cell actin maintenance and hearing.

    Evidence Hair bundle proteomics, CRISPR/Cas9 Xirp2-null mice, ABR, and SEM/TEM ultrastructure, replicated across two studies

    PMID:25653358 PMID:25772365

    Open questions at the time
    • Differential targeting determinants of short vs long isoforms unmapped
    • Whether degeneration reflects failed repair vs structural support unclear at this stage
    • Direct stereocilia binding partners not identified
  6. 2016 Medium

    Identified filamin C as a stabilization client, linking XIRP2 actin-crosslinking to maintenance of specific sarcomeric components in nascent myofibrils.

    Evidence FRAP in Xirp1/Xirp2-deficient cardiomyocytes and live zebrafish imaging

    PMID:27206985

    Open questions at the time
    • Single lab
    • Direct vs indirect stabilization of FLNc not separated
    • Redundancy with XIRP1 not fully partitioned
  7. 2018 High

    Mechanistically linked the ICD/cytoskeletal role to electrophysiology, showing XIRP2 associates with cardiac ion channels and is required for normal conduction.

    Evidence Xirp2 knockout mouse ECG, whole-cell patch-clamp, and co-IP with Nav1.5 and Kv1.5

    PMID:29306897

    Open questions at the time
    • Whether XIRP2 directly traffics channels or stabilizes them is unresolved
    • Co-IP not validated reciprocally
    • Direct vs ICD-architecture-mediated channel effects not separated
  8. 2018 Medium

    Revealed that stereocilia cross-linkers including XIRP2 turn over continuously despite a stable actin core, framing repair as ongoing compositional remodeling.

    Evidence Transgenic EGFP-fascin-2 mouse imaging with endogenous cross-linker comparison

    PMID:29874122

    Open questions at the time
    • XIRP2 turnover inferred by displacement, not directly tracked
    • Regulation of turnover unknown
    • Single lab
  9. 2023 High

    Defined the repair mechanism: a C-terminal mechanosensor domain confers force-dependent recruitment of XIRP2 to actin damage sites to drive monomeric actin enrichment and repair.

    Evidence Xirp2-null mice with traumatic noise, phalloidin gap and monomeric actin assays, and fibroblast stretch with domain deletion mapping

    PMID:37294664

    Open questions at the time
    • Molecular nature of the force-sensing conformational change unresolved
    • How XIRP2 promotes actin monomer delivery mechanistically unclear
    • Whether the cardiac functions use the same mechanosensor domain untested
  10. 2015 Medium

    Provided human disease relevance, linking truncating XIRP2 mutations to dilated cardiomyopathy via the same ICD defect seen in mice.

    Evidence Whole exome sequencing of a DCM child with cardiac histomorphology and cross-species phenotype comparison

    PMID:26656454

    Open questions at the time
    • Single case
    • Modifier vs primary causative role not established
    • Functional consequence of the specific truncations not tested in vitro
  11. 2024 Medium

    Implicated XIRP2 in cancer-cell ion homeostasis and chemosensitivity, suggesting a role beyond cytoskeletal architecture.

    Evidence XIRP2 knockdown in hepatocellular carcinoma cells with drug sensitivity, ion measurements, and protein stability assays

    PMID:39194571

    Open questions at the time
    • Mechanism linking XIRP2 to zinc/calcium regulation undefined
    • Single lab with limited mechanistic depth
    • Relationship to the cytoskeletal/ion-channel roles unclear

Open questions

Synthesis pass · forward-looking unresolved questions
  • How XIRP2's single actin-crosslinking and mechanosensing biochemistry is deployed across such divergent contexts — cardiac ICD/channel organization, sarcomeric stabilization, stereocilia repair, and ion homeostasis — remains unresolved.
  • No structure of the actin-binding repeats or mechanosensor domain
  • Unified model relating tissue-specific isoforms to function lacking
  • Direct vs scaffolding role in ion channel surface expression not established

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0005198 structural molecule activity 2 GO:0008092 cytoskeletal protein binding 2 GO:0140299 molecular sensor activity 1
Localization
GO:0005856 cytoskeleton 3 GO:0005829 cytosol 2
Pathway
R-HSA-9709957 Sensory Perception 3 R-HSA-397014 Muscle contraction 2 R-HSA-162582 Signal Transduction 1
Partners
ACTN2FLNCKCNA5SCN5A
Complex memberships
intercalated discstereocilia actin core

Evidence

Reading pass · 12 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2004 The Xin repeats of XIRP2 (called 'Xin repeat protein 2') define a novel actin-binding motif: in vitro co-sedimentation assays with skeletal muscle actin demonstrated that XIRP2's 28 Xin repeats directly bind actin filaments and can arrange microfilaments into networks that sediment upon low-speed centrifugation. Transfection experiments confirmed that the repeats bind and stabilize the actin-based cytoskeleton in cultured cells. In vitro co-sedimentation assay with skeletal muscle actin; transfection of repeat constructs into cultured cells Journal of cell science High 15454575
2006 XIRP2 (mouse ortholog myomaxin/CMYA3) is a direct transcriptional target of MEF2A and localizes to the Z-disc/costameric region in striated muscle. The myomaxin protein (mouse XIRP2 ortholog) interacts with the sarcomeric Z-disc protein alpha-actinin-2. MEF2A knockout mouse expression analysis (microarray); protein interaction demonstrated by co-immunoprecipitation and/or pulldown; immunolocalization in muscle The Journal of biological chemistry Medium 17046827
2010 XIRP2 expression in the heart is induced by angiotensin II (Ang II) signaling through MEF2A transcriptional activity. Ang II activates XIRP2 promoter activity via MEF2A, placing XIRP2 downstream of the Ang II–MEF2A pathway. Xirp2 hypomorphic mice (with markedly reduced cardiac Xirp2) developed spontaneous cardiac hypertrophy and increased beta-myosin heavy chain expression, and showed attenuated hypertrophic remodeling, reduced fibrosis, and reduced apoptosis after chronic Ang II infusion. Xirp2 promoter characterization; MEF2A activity assay; Xirp2 hypomorphic mouse model with Ang II infusion; histological and molecular readouts Circulation research High 20093629
2012 Mouse Xinβ (mXinβ/XIRP2) is required for the postnatal maturation of intercalated discs (ICDs) in the heart. In mXinβ-null mice, intercellular junction components fail to redistribute from the lateral membrane to the termini of cardiomyocytes during postnatal ICD formation (defect apparent at postnatal day 16.5). mXinβ is uniquely upregulated during the redistribution phase and preferentially associates with forming ICDs by immunofluorescence and subcellular fractionation. mXinβ knockout mouse; quantitative Western blot; immunofluorescence; subcellular fractionation; comparison with mXinα-null and double-knockout hearts Developmental biology High 23261932
2015 A short splice form of XIRP2 (lacking the Xin repeats, also called XEPLIN) is expressed in inner ear hair cells where it colocalizes with actin-rich stereocilia, the cuticular plate, and the circumferential actin belt. CRISPR/Cas9-mediated Xirp2-null mice exhibit high-frequency hearing loss and stereocilia degeneration, demonstrating that XIRP2 is required for long-term maintenance of hair cell stereocilia morphology and hearing function. Mass spectrometry of hair bundle proteome; CRISPR/Cas9 knockout mouse; auditory brainstem response (hearing function); scanning electron microscopy of stereocilia The Journal of neuroscience High 25653358
2015 Different isoforms of XIRP2 are differentially localized in hair cells: short splice forms (XEPLIN) are targeted preferentially to stereocilia, whereas two long isoforms containing the XIN-repeat domain are present in both stereocilia and cuticular plates. Xirp2-null mice develop normal stereocilia bundles that degenerate over time, with loss of stereocilia, long membranous protrusions from apical surfaces, and disorganization of paracrystalline actin filaments at the ultrastructural level. Isoform-specific immunolocalization; Xirp2 knockout mouse; scanning electron microscopy; transmission electron microscopy of actin ultrastructure Cell reports High 25772365
2016 Both Xirp1 and Xirp2 (Xin actin-binding repeat-containing proteins) stabilize filamin C (FLNc) selectively in premyofibrils of cardiomyocytes. Analysis of cardiomyocytes from Xirp1- and Xirp2-deficient animals showed reduced FLNc stability at premyofibrils, as assessed by fluorescence recovery after photobleaching. FRAP (fluorescence recovery after photobleaching) in cardiomyocytes from Xirp1/Xirp2 knockout animals; live zebrafish imaging Human molecular genetics Medium 27206985
2018 Xirp2 is required for normal cardiac conduction. Xirp2-knockout mouse hearts exhibit prolonged PR and QT intervals, slow conduction velocity, atrioventricular conduction block, and abnormal infranodal ventricular conduction. Whole-cell patch-clamp of Xirp2-/- cardiomyocytes detected altered ionic currents. Co-immunoprecipitation demonstrated association of Xirp2 with Nav1.5 and Kv1.5 ion channel subunits, consistent with a role for Xirp2 in governing ion channel surface expression and cardiac conduction. Xirp2 knockout mouse; electrocardiography; whole-cell patch-clamp; co-immunoprecipitation with Nav1.5 and Kv1.5 Journal of the American Heart Association High 29306897
2018 In stereocilia, actin cross-linkers including XIRP2 undergo continuous turnover even though the actin filament core is highly stable, demonstrating that ongoing remodeling of cross-linker composition is a mechanism for stereocilia homeostasis and repair. Transgenic mouse expressing EGFP-fascin-2 to monitor cross-linker dynamics; comparison with endogenous XIRP2, espin, and plastin-1 localization Molecular biology of the cell Medium 29874122
2023 XIRP2 is required for repair of noise-induced damage to stereocilia F-actin cores. XIRP2 facilitates enrichment of monomeric γ-actin at damage gaps in auditory hair cells. Recruitment of XIRP2 to stereocilia gaps and to stress fiber strain sites in fibroblasts is force-dependent and is mediated by a novel mechanosensor domain located in the C-terminus of XIRP2. Xirp2-null mice subjected to traumatic noise; phalloidin gap assay; monomeric actin immunostaining; fibroblast stretch assay for mechanosensor domain mapping; domain deletion/mutagenesis experiments eLife High 37294664
2015 Compound heterozygous truncating mutations in XIRP2 were identified in a child with sporadic dilated cardiomyopathy. Histomorphological analysis of explanted heart tissue showed misregistration, mislocalization, and shortening of intercalated discs, findings similar to those in Xirp2-/- mice, implicating XIRP2 as a modifier gene in DCM pathogenesis through disruption of ICD integrity. Whole exome sequencing; cardiac histomorphology of explanted heart; comparison with Xirp2-/- mouse phenotype Journal of the American Heart Association Medium 26656454
2024 Inhibition of XIRP2 in hepatocellular carcinoma cells results in increased sensitivity to oxaliplatin through elevation of zinc ions and calcium ion overload, identifying a functional role for XIRP2 in ion homeostasis. The XIRP2 mutation was found to enhance XIRP2 protein stability without affecting mRNA levels. XIRP2 knockdown in HCC cells; CCK-8 drug sensitivity assay; protein stability experiments; ion (zinc/calcium) measurements Biology Medium 39194571

Source papers

Stage 0 corpus · 61 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2020 Genomic landscape of metastatic breast cancer identifies preferentially dysregulated pathways and targets. The Journal of clinical investigation 95 32657779
2004 Xin repeats define a novel actin-binding motif. Journal of cell science 83 15454575
2014 Genome-wide association and pathway analysis of feed efficiency in pigs reveal candidate genes and pathways for residual feed intake. Frontiers in genetics 76 25250046
2016 Filamin C is a highly dynamic protein associated with fast repair of myofibrillar microdamage. Human molecular genetics 72 27206985
2020 EP300 mutation is associated with tumor mutation burden and promotes antitumor immunity in bladder cancer patients. Aging 64 32012118
2016 Parkinson's disease: SNCA-, PARK2-, and LRRK2- targeting microRNAs elevated in cingulate gyrus. Parkinsonism & related disorders 63 27717584
2010 Modulation of angiotensin II-mediated cardiac remodeling by the MEF2A target gene Xirp2. Circulation research 61 20093629
2013 Differential proteomic analysis of abnormal intramyoplasmic aggregates in desminopathy. Journal of proteomics 60 23639843
2022 Single-cell and spatial transcriptomics of the infarcted heart define the dynamic onset of the border zone in response to mechanical destabilization. Nature cardiovascular research 54 39086770
2010 Genome-wide association study identifies genes that may contribute to risk for developing heroin addiction. Psychiatric genetics 54 20520587
2006 Myomaxin is a novel transcriptional target of MEF2A that encodes a Xin-related alpha-actinin-interacting protein. The Journal of biological chemistry 50 17046827
2015 A short splice form of Xin-actin binding repeat containing 2 (XIRP2) lacking the Xin repeats is required for maintenance of stereocilia morphology and hearing function. The Journal of neuroscience : the official journal of the Society for Neuroscience 43 25653358
2010 Cardioprotective effects of a selective B(2) receptor agonist of bradykinin post-acute myocardial infarct. American journal of hypertension 41 20186129
2015 XIRP2, an actin-binding protein essential for inner ear hair-cell stereocilia. Cell reports 40 25772365
2006 SLC25A12 and CMYA3 gene variants are not associated with autism in the IMGSAC multiplex family sample. European journal of human genetics : EJHG 38 16205742
2021 Right versus left ventricular remodeling in heart failure due to chronic volume overload. Scientific reports 36 34429479
2018 Transcriptome Analysis of Cardiac Hypertrophic Growth in MYBPC3-Null Mice Suggests Early Responders in Hypertrophic Remodeling. Frontiers in physiology 34 30410445
2014 New insights into the roles of Xin repeat-containing proteins in cardiac development, function, and disease. International review of cell and molecular biology 32 24725425
2012 The Xin repeat-containing protein, mXinβ, initiates the maturation of the intercalated discs during postnatal heart development. Developmental biology 28 23261932
2012 Patient-specific protein aggregates in myofibrillar myopathies: laser microdissection and differential proteomics for identification of plaque components. Proteomics 26 23044792
2005 Structure, Expression, and Function of a Novel Intercalated Disc Protein, Xin. Journal of medical sciences (Taipei, Taiwan) 26 16708114
2022 Chronic isoprenaline/phenylephrine vs. exclusive isoprenaline stimulation in mice: critical contribution of alpha1-adrenoceptors to early cardiac stress responses. Basic research in cardiology 25 35286475
2015 Identification of quantitative trait loci influencing inflammation-mediated alveolar bone loss: insights into polygenic inheritance of host-biofilm disequilibria in periodontitis. Journal of periodontal research 22 26126603
2015 Exome Sequencing Identifies Pathogenic and Modifier Mutations in a Child With Sporadic Dilated Cardiomyopathy. Journal of the American Heart Association 22 26656454
2018 Critical Roles of Xirp Proteins in Cardiac Conduction and Their Rare Variants Identified in Sudden Unexplained Nocturnal Death Syndrome and Brugada Syndrome in Chinese Han Population. Journal of the American Heart Association 21 29306897
2018 The stable actin core of mechanosensory stereocilia features continuous turnover of actin cross-linkers. Molecular biology of the cell 20 29874122
2009 The candidate gene XIRP2 at a quantitative gene locus on equine chromosome 18 associated with osteochondrosis in fetlock and hock joints of South German Coldblood horses. The Journal of heredity 16 19304740
2023 Identification of copy number variation in Tibetan sheep using whole genome resequencing reveals evidence of genomic selection. BMC genomics 15 37726692
2021 Adaptive versus maladaptive cardiac remodelling in response to sustained β-adrenergic stimulation in a new 'ISO on/off model'. PloS one 14 34138844
2020 Identifying modifier genes for hypertrophic cardiomyopathy. Journal of molecular and cellular cardiology 14 32470469
2020 Transformation of Mature Osteoblasts into Bone Lining Cells and RNA Sequencing-Based Transcriptome Profiling of Mouse Bone during Mechanical Unloading. Endocrinology and metabolism (Seoul, Korea) 14 32615730
2006 A novel gene (Cmya3) induced in the heart by angiotensin II-dependent but not salt-dependent hypertension in mice. American journal of hypertension 14 16500513
2013 Intercalated disc protein, mXinα, suppresses p120-catenin-induced branching phenotype via its interactions with p120-catenin and cortactin. Archives of biochemistry and biophysics 13 23296090
2023 Repair of noise-induced damage to stereocilia F-actin cores is facilitated by XIRP2 and its novel mechanosensor domain. eLife 12 37294664
2018 Genetic Variants Influencing Plasma Renin Activity in Hypertensive Patients From the PEAR Study (Pharmacogenomic Evaluation of Antihypertensive Responses). Circulation. Genomic and precision medicine 12 29650764
2022 Methylome analysis of ALS patients and presymptomatic mutation carriers in blood cells. Neurobiology of aging 11 35537341
2016 Sarcomeric lesions and remodeling proximal to intercalated disks in overload-induced cardiac hypertrophy. Experimental cell research 11 27639425
2022 Whole exome sequencing of high-risk neuroblastoma identifies novel non-synonymous variants. PloS one 10 36037157
2018 Germline but not somatic de novo mutations are common in human congenital diaphragmatic hernia. Birth defects research 10 29570242
2024 Ca2+/calmodulin-dependent kinase IIδC-induced chronic heart failure does not depend on sarcoplasmic reticulum Ca2+ leak. ESC heart failure 9 38616546
2024 Interleukin 11 therapy causes acute left ventricular dysfunction. Cardiovascular research 8 39383190
2022 A genomic mutation spectrum of collecting duct carcinoma in the Chinese population. BMC medical genomics 8 34980126
2022 RNA-Seq analysis reveals sex-dependent transcriptomic profiles of human subacromial bursa stratified by tear etiology. Journal of orthopaedic research : official publication of the Orthopaedic Research Society 8 35266580
2021 Genome-wide analysis of the mouse LIM gene family reveals its roles in regulating pathological cardiac hypertrophy. FEBS letters 8 34328660
2022 Mutation burden analysis of six common mental disorders in African Americans by whole genome sequencing. Human molecular genetics 6 35642741
2023 Effects of Purine Metabolism-Related LINC01671 on Tumor Heterogeneity in Kidney Renal Clear Cell Carcinoma. Frontiers in bioscience (Landmark edition) 5 38179759
2018 Polymorphisms in selected genes and analysis of their relationship with osteochondrosis in Polish sport horse breeds. Animal genetics 5 30152531
2025 Genome-Wide Association Study of Exercise Addiction Among Elite Wrestlers. Brain sciences 4 40002435
2014 CCDD Phenotype Associated with a Small Chromosome 2 Deletion. Seminars in ophthalmology 4 24475916
2024 Identification of Clinical Value and Biological Effects of XIRP2 Mutation in Hepatocellular Carcinoma. Biology 3 39194571
2025 Identification of Potential Intervention Targets Involved in Prior Exercise that Attenuates Peripheral Neuropathic Pain by Integrating Transcriptome and Whole-genome Bisulfite Sequencing Analyses. Molecular neurobiology 2 39825163
2024 Somatic mutations that affect early genetic progression and immune microenvironment in gastric carcinoma. Pathology, research and practice 2 38663178
2025 Whole-exome sequencing reveals the mutational landscape of head and neck lymphoepithelioma-like carcinoma. Oral oncology 1 40976151
2025 R405W Desmin Knock-In Mice Highlight Alterations of Mitochondria, Protein Quality Control and Myofibrils in Myofibrillar Myopathy. Journal of cachexia, sarcopenia and muscle 1 41165044
2023 A Rare Case of Penoscrotal Webbing and Extensive Hernias: An Anatomical Report With Genetic Insights. Cureus 1 38021525
2026 Genomic Signatures Underlying Environmental Adaptation and Reproductive Traits in the Tibetan Pig. Animals : an open access journal from MDPI 0 41681490
2025 Early downregulation of hair cell (HC)-specific genes in the vestibular sensory epithelium during chronic ototoxicity. Journal of biomedical science 0 40908485
2025 Transcriptome analysis of postnatal mouse cardiac tissue growth and development. Yi chuan = Hereditas 0 41139466
2025 Nuclear Ca2+-Calmodulin signaling in cardiac myocytes reduces catecholamine-evoked protein translation and prevents hypertrophy. Cell calcium 0 41380480
2024 Comparative Proteomic and Phosphoproteomic Analyses Reveal Molecular Signatures of Myocardial Infarction and Transverse Aortic Constriction in Aged Mouse Models. Cardiology research and practice 0 39502510
2024 Multi-ancestry genome-wide association study reveals novel genetic signals for lung function decline. medRxiv : the preprint server for health sciences 0 39649580

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