Affinage

ORAI2

Protein orai-2 · UniProt Q96SN7

Length
254 aa
Mass
28.6 kDa
Annotated
2026-06-10
27 papers in source corpus 18 papers cited in narrative 18 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

ORAI2 is a pore-forming subunit of store-operated Ca2+ release-activated Ca2+ (CRAC) channels that, together with STIM1, conducts Ca2+-selective currents upon store depletion, with intrinsic conductance efficacy ranking ORAI1 > ORAI2 > ORAI3 (PMID:16807233). In most cell types ORAI2 assembles into hetero-oligomeric (likely hetero-tetrameric) channels with ORAI1, where it acts as a negative rheostat: overexpression lowers and loss-of-function raises store-operated Ca2+ entry (SOCE), as demonstrated by single-molecule coupling in chondrocytes and reciprocal knockout phenotypes in T cells (PMID:25769459, PMID:28294127). This attenuating role is conserved across mast cells, astrocytes, and neuroglioma, where ORAI2 deletion or knockdown augments CRAC currents and amplifies downstream Ca2+-dependent outputs including mast cell degranulation and anaphylaxis, astrocytic PGE2 production, and amyloid-beta handling (PMID:29604961, PMID:35506586, PMID:32722509). A second ORAI2 splice variant (ORAI2S) displays strong Ca2+-dependent inactivation and a dominant-negative effect on channel assembly, providing an additional layer of conductance tuning (PMID:17463004). In specific cellular contexts ORAI2 instead serves as the primary positive driver of SOCE: it is the dominant pore subunit in neutrophils, where its loss decreases Ca2+ influx through a KCa3.1-dependent membrane-potential mechanism, and it sustains capacitative entry in neurons, where Orai2 deficiency protects against ischemic neuronal death (PMID:32929002, PMID:31551038). ORAI2-mediated SOCE feeds defined signaling axes that govern proliferation, migration, and pathology — activating PI3K/Akt and FAK/MAPK/ERK to promote gastric cancer growth and metastasis, and an ORAI2/JNK/NFAT1 axis driving TGF-beta1-mediated salivary gland fibrosis (PMID:33310726, PMID:39384103). ORAI2 expression is itself regulated post-transcriptionally by NSUN2-mediated m5C modification of its mRNA, read by YBX1 to enhance transcript stability (PMID:37130916), and is cell-cycle-dependent, peaking at G2/M to restrain SOCE (PMID:25748572).

Mechanistic history

Synthesis pass · year-by-year structured walk · 11 steps
  1. 2006 High

    Established that ORAI2 is not merely an ORAI1 homolog but an autonomous pore-forming CRAC channel subunit, answering whether it could conduct store-operated Ca2+ current at all.

    Evidence Co-transfection of ORAI2 with STIM1 in HEK293 cells and electrophysiological recording of Ca2+-selective store-operated currents

    PMID:16807233

    Open questions at the time
    • Did not address ORAI2 behavior in heteromeric channels with ORAI1
    • Quantitative efficacy ranking measured only in heterologous overexpression
  2. 2007 High

    Revealed splice-variant and cell-background control of ORAI2 channel function, showing ORAI2S can act dominant-negatively on CRAC assembly and adding a tuning mechanism beyond simple expression level.

    Evidence Cloning of murine ORAI2L/ORAI2S splice variants, expression with STIM1 in HEK293 and RBL 2H3 cells, whole-cell patch-clamp

    PMID:17463004

    Open questions at the time
    • Functional difference between HEK293 and RBL 2H3 not mechanistically explained
    • Physiological relevance of ORAI2S dominant-negative effect in native cells not established
  3. 2014 Medium

    Proposed a non-canonical localization of ORAI2 on secretory granules and a role in antigen-specific Ca2+ mobilization, distinct from a purely plasma-membrane channel function.

    Evidence Subcellular fractionation, immunofluorescence, siRNA knockdown, Ca2+ imaging and exocytosis assay in RBL-2H3 mast cells

    PMID:25044118

    Open questions at the time
    • Granular localization shown by fractionation/immunofluorescence only, no reconstitution of granule channel activity
    • Single cell line; not confirmed in primary mast cells
  4. 2015 High

    Defined ORAI2 as a negative modulator within ORAI1-ORAI2 heteromeric channels, answering how it shapes SOCE magnitude rather than simply adding conductance.

    Evidence Single-molecule TIRF imaging of ORAI1-ORAI2 coupling plus reciprocal knockdown/overexpression and Ca2+ imaging in chondrocytes

    PMID:25769459

    Open questions at the time
    • Exact stoichiometry of the hetero-oligomer not resolved
    • Whether attenuation arises from altered gating or reduced ORAI1 incorporation not determined
  5. 2015 Medium

    Linked ORAI2 expression and SOCE to cell-state transitions and proliferation, beginning to connect channel function to physiological outcomes.

    Evidence qPCR, Western blot, Ca2+ imaging and siRNA knockdown in pulmonary arterial smooth muscle and brain capillary endothelial cell-cycle models

    PMID:25673771 PMID:25748572

    Open questions at the time
    • Pathway placement largely correlative without epistasis
    • Cell-cycle-dependent upregulation mechanism at G2/M not defined
  6. 2017 High

    Provided definitive in vivo genetic proof that ORAI2 attenuates CRAC function in immune cells, with reciprocal SOCE phenotypes for Orai1 vs Orai2 deletion and physiological consequences for T cell and mast cell biology.

    Evidence Orai1, Orai2 single and double knockout mice with CRAC electrophysiology, Ca2+ imaging, Co-IP/FRET, and in vivo immune and anaphylaxis models

    PMID:28294127 PMID:29604961

    Open questions at the time
    • Molecular basis of the attenuating effect within native heteromers not resolved
    • Tissue-specific expression ratios governing the degree of attenuation not quantified
  7. 2019 High

    Showed ORAI2 can be the dominant positive mediator of SOCE in specific lineages, overturning a uniform attenuator model; established neuronal and dendritic-cell roles and a regulatory link to LRRK2.

    Evidence Orai2 knockout neurons and mice with Ca2+ imaging and stroke models; LRRK2-KO/inhibitor with isoform-specific siRNA and migration assays

    PMID:31166814 PMID:31551038

    Open questions at the time
    • Why ORAI2 is positive in neurons but attenuating elsewhere not mechanistically explained
    • Mechanism by which LRRK2 kinase activity selectively suppresses ORAI2 expression unknown
  8. 2020 High

    Resolved the context-dependence by identifying a KCa3.1-dependent membrane-potential mechanism that makes ORAI2 the primary positive SOCE driver in neutrophils, and connected ORAI2 SOCE to oncogenic PI3K/Akt and FAK/MAPK/ERK signaling in cancer.

    Evidence Neutrophil knockout mice with patch-clamp and membrane-potential measurements; gain/loss-of-function in gastric cancer lines and xenografts with signaling readouts

    PMID:27865925 PMID:32929002 PMID:33310726

    Open questions at the time
    • How KCa3.1 coupling differs between neutrophils and attenuator cell types not defined
    • Direct biochemical link between ORAI2 channel activity and PI3K/FAK activation not established
  9. 2021 High

    Distinguished ORAI2 biophysically from its homologs and extended its negative-regulatory role to amyloid-beta production, refining isoform-specific channel properties.

    Evidence Whole-cell patch-clamp with pH manipulation and ORAI1-ORAI3 chimeras; siRNA/overexpression with Abeta42 ELISA in H4-APPswe cells

    PMID:32722509 PMID:34877682

    Open questions at the time
    • Structural determinant of ORAI2 pH-insensitive inactivation not mapped to specific residues
    • Abeta findings limited to a single neuroglioma cell line
  10. 2022 Medium

    Demonstrated ORAI2 restrains inflammatory and fibrotic Ca2+-dependent programs, linking it to CNS PGE2 production and to a JNK/NFAT1/TGF-beta1 fibrogenic axis with therapeutic relevance.

    Evidence Astrocyte siRNA/KO with Ca2+ imaging, qPCR/Western and PGE2 ELISA; irradiated salivary gland model with SOCE/NFAT1 inhibition and ORAI2 KO

    PMID:35506586 PMID:39384103

    Open questions at the time
    • Whether ORAI2-PGE2 and ORAI2-fibrosis effects are direct channel functions or secondary to ORAI1 upregulation not separated
    • NFAT1 activation mechanism downstream of ORAI2 in fibroblasts not detailed
  11. 2023 Medium

    Identified the upstream post-transcriptional control of ORAI2 abundance, explaining how its expression is tuned in disease via an NSUN2/m5C/YBX1 axis.

    Evidence RNA bisulfite sequencing, RIP for NSUN2/YBX1, mRNA stability assays, E2F1 ChIP and metastasis assays in gastric cancer

    PMID:37130916

    Open questions at the time
    • Whether m5C regulation of ORAI2 operates outside gastric cancer not tested
    • Single lab; YBX1-dependent stabilization not reconstituted in vitro

Open questions

Synthesis pass · forward-looking unresolved questions
  • The molecular determinant that switches ORAI2 between attenuator (T cells, mast cells, astrocytes) and primary positive conductor (neutrophils, neurons) remains unresolved.
  • No structural model of native ORAI1-ORAI2 heteromer stoichiometry across cell types
  • Relative ORAI1/ORAI2/STIM expression ratios that dictate sign of the effect not systematically mapped

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0005215 transporter activity 5
Localization
GO:0005886 plasma membrane 3 GO:0031410 cytoplasmic vesicle 1
Pathway
R-HSA-1643685 Disease 4 R-HSA-162582 Signal Transduction 3 R-HSA-168256 Immune System 3
Partners
ORAI1STIM1STIM2
Complex memberships
ORAI1-ORAI2 heteromeric CRAC channel

Evidence

Reading pass · 18 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2006 Co-expression of ORAI2 with STIM1 in HEK293 cells produces large store-operated Ca2+-selective currents (CRAC currents), demonstrating that ORAI2 can function as a pore-forming subunit of store-operated Ca2+ channels together with STIM1; efficacy is in the order ORAI1 > ORAI2 > ORAI3. Co-transfection of HEK293 cells with STIM1 and ORAI2 followed by electrophysiological recording of Ca2+-selective store-operated currents and Ca2+ entry measurements The Journal of biological chemistry High 16807233
2007 Murine ORAI2 produces two splice variants (ORAI2L and ORAI2S) that both form functional CRAC channels when co-expressed with STIM1 in HEK293 cells, but not in RBL 2H3 cells, indicating cell-background dependence; ORAI2S shows strong Ca2+-dependent inactivation and exerts a dominant-negative effect on CRAC channel formation when co-expressed with STIM1 and ORAI1. Cloning of splice variants, Northern blot tissue expression, co-expression with STIM1 in HEK293 and RBL 2H3 cells, whole-cell patch-clamp electrophysiology The Journal of biological chemistry High 17463004
2015 ORAI2 forms a hetero-oligomeric (likely hetero-tetramer) complex with ORAI1 in chondrocyte cells, as demonstrated by single-molecule TIRF imaging showing direct coupling; overexpression of ORAI2 decreases SOCE while knockdown increases SOCE, indicating ORAI2 reduces ORAI1 function within the heteromeric channel. siRNA knockdown and overexpression, fluorescence Ca2+ imaging, single-molecule total internal reflection fluorescence (TIRF) microscopy showing ORAI1-ORAI2 direct coupling Cell calcium High 25769459
2017 ORAI2 forms heteromeric CRAC channels with ORAI1 in mouse T cells and attenuates CRAC channel function; deletion of Orai2 increases SOCE, while deletion of Orai1 reduces SOCE; combined deletion of Orai1 and Orai2 abolishes SOCE and strongly impairs T cell function in vitro and in vivo, including antiviral responses and T cell-mediated autoimmunity. Genetic deletion (Orai1-/-, Orai2-/-, Orai1/Orai2 double-KO mice), Ca2+ imaging, electrophysiology (CRAC current), co-immunoprecipitation/FRET for heteromeric channel formation, in vivo mouse models of colitis and graft-versus-host disease Nature communications High 28294127
2017 Deletion of Orai2 in murine peritoneal mast cells augments endogenous CRAC currents, enhances Ca2+ rise triggered by IgE receptor or MAS-related GPCR stimulation, increases Ca2+-dependent degranulation, and intensifies mast cell-mediated anaphylaxis in vivo, demonstrating that Orai2 limits CRAC channel activity in mast cells. Orai2 gene knockout mouse, whole-cell patch-clamp (CRAC current density), Ca2+ imaging, degranulation assay, in vivo anaphylaxis model Cell calcium High 29604961
2014 ORAI2 is localized on secretory granules (not exclusively plasma membrane) in RBL-2H3 mast cells; knockdown of Orai2 attenuates antigen-stimulated Ca2+ release from Ca2+ stores and inhibits exocytotic release, but does not affect thapsigargin-induced Ca2+ release, suggesting a role in antigen-specific Ca2+ mobilization from granules. Subcellular fractionation and immunofluorescence localization, siRNA knockdown, Ca2+ imaging, exocytosis assay in RBL-2H3 cells Biochemical and biophysical research communications Medium 25044118
2015 In proliferating (vs. contractile) pulmonary arterial smooth muscle cells, STIM2 and ORAI2 are upregulated and associated with increased store-operated Ca2+ entry (SOCE), whereas contractile-state cells show increased voltage-dependent Ca2+ entry; knockdown/functional data indicate STIM2 and Orai2 upregulation drives phenotypic transition to a proliferative state. Quantitative PCR, Western blot, Ca2+ imaging (SOCE assay), siRNA knockdown in PASMC phenotype switch model American journal of physiology. Cell physiology Medium 25673771
2019 ORAI2 is a central mediator of neuronal capacitative Ca2+ entry; Orai2-deficient neurons show significantly diminished Ca2+ signals induced by store depletion or oxygen/glucose deprivation, and Orai2-deficient mice are protected from ischemic neuronal death during acute stroke and ischemia/reperfusion. Genetic knockout mouse, Ca2+ imaging in primary neurons (store depletion and OGD), middle cerebral artery occlusion experimental stroke model Stroke High 31551038
2015 ORAI2 expression is specifically upregulated at the G2/M phase of the cell cycle in brain capillary endothelial cells; at this phase, increased ORAI2 negatively modulates SOCE activity, and knockdown of ORAI2 at G2/M removes this SOCE decrease and partly attenuates cell proliferation. Cell synchronization by double thymidine blockage, qPCR and Western blot, siRNA knockdown, Ca2+ imaging (SOCE assay), cell proliferation assay Biochemical and biophysical research communications Medium 25748572
2016 ORAI2 contributes to SOCE in HL60 leukemia cells; Orai2 silencing significantly attenuates thapsigargin-induced SOCE, and combined silencing of Orai1 and Orai2 nearly abolishes SOCE; knockdown of Orai1 and Orai2 impairs HL60 cell migration in vitro, associated with impaired FAK tyrosine phosphorylation. siRNA knockdown, fluorescence Ca2+ imaging, transwell migration assay, Western blot for FAK phosphorylation Biochimica et biophysica acta. Molecular cell research Medium 27865925
2020 ORAI2 mediates store-operated Ca2+ entry and promotes gastric cancer cell proliferation, migration, tumor formation and metastasis; mechanistically, ORAI2-mediated SOC activity activates PI3K/Akt signaling and enhances FAK-mediated MAPK/ERK activation, promoting focal adhesion disassembly at the rear edge of migrating cells. Gain- and loss-of-function (overexpression and siRNA/shRNA knockdown) in gastric cancer cell lines and xenograft mouse models; Ca2+ imaging, Western blot for PI3K/Akt and FAK/MAPK/ERK, focal adhesion assays Cancer research Medium 33310726
2020 ORAI2 is the primary pore subunit mediating capacitative Ca2+ entry in murine neutrophils; ORAI2-deficient neutrophils show decreased (not increased) Ca2+ influx, contrasting with enhanced SOCE seen in other immune cell types lacking ORAI2; this decreased SOCE in ORAI2-deficient neutrophils is correlated with altered KCa3.1-mediated membrane potential regulation. Genetic KO mouse (Orai1-/-, Orai2-/-, double KO), Ca2+ imaging, patch-clamp, membrane potential measurements, neutrophil functional assays (phagocytosis, degranulation, ROS, leukotriene), in vivo staphylococcal infection model Proceedings of the National Academy of Sciences of the United States of America High 32929002
2019 LRRK2 kinase activity negatively regulates ORAI2 expression in dendritic cells; LRRK2 deficiency increases ORAI2 expression (but not ORAI1 or ORAI3) and enhances DC migration; silencing ORAI2 markedly decreases DC migration without affecting maturation markers, whereas ORAI1 silencing affects both migration and maturation markers. Genetic KO (LRRK2-/- mice), pharmacological LRRK2 kinase inhibition, siRNA knockdown of ORAI1/2/3, Ca2+ imaging, transwell migration assay, flow cytometry for maturation markers FASEB journal Medium 31166814
2021 ORAI2 amplitude of CRAC current (ICRAC) is sensitive to changes in intracellular pH similarly to ORAI1, but unlike ORAI1, fast Ca2+-dependent inactivation kinetics of ORAI2 are unaffected by acidic intracellular pH; ORAI3-mediated ICRAC shows no pH dependence. Whole-cell patch-clamp of HEK293T cells heterologously expressing ORAI isoforms with STIM1, intracellular pH manipulation, Orai1-Orai3 chimera domain-swap analysis The Journal of physiology High 34877682
2022 ORAI2 negatively modulates SOCE in astrocytes; Orai2 knockdown or knockout increases SOCE and Orai1 expression, and augments production of prostaglandin E2 (via upregulated Ptges and Ptgs2) in TNFα/IL1α-stimulated astrocytes, demonstrating that ORAI2 restrains inflammatory PGE2 production in the CNS. siRNA knockdown and genetic KO (Orai2-KO) in primary astrocytes, Ca2+ imaging, qPCR and Western blot for Ptges/Ptgs2/Orai1, PGE2 ELISA Glia Medium 35506586
2023 NSUN2-mediated m5C modification of ORAI2 mRNA increases its stability and expression; YBX1 acts as the 'reader' of this m5C modification site; upstream, fatty acids from omental adipocytes activate the AMPK pathway to upregulate E2F1, which promotes NSUN2 transcription, thereby driving ORAI2 expression and gastric cancer peritoneal metastasis. RNA bisulfite sequencing (m5C mapping), RIP (RNA immunoprecipitation) for NSUN2 and YBX1, mRNA stability assay, ChIP for E2F1 on NSUN2 promoter, functional colonization/metastasis assays Oncogene Medium 37130916
2024 ORAI2 mediates a SOCE-dependent signaling axis (ORAI2/JNK/NFAT1) that drives TGF-β1-mediated fibrogenesis in irradiated salivary glands; inhibition of SOCE reduces fibrosis in an ORAI2-dependent manner, and pharmacological inhibition of NFAT1 restores saliva flow to ~85% of normal in irradiated mice. RNA sequencing of irradiated mouse salivary glands, SOCE inhibition (SKF96365, YM58483), siRNA/KO of ORAI2, Western blot for JNK/NFAT1/TGF-β1, in vivo irradiation model with pharmacological NFAT1 inhibition International journal of radiation oncology, biology, physics Medium 39384103
2021 ORAI2 downregulation in human neuroglioma H4 cells significantly increases SOCE amplitude; in Aβ-secreting H4-APPswe cells, ORAI2 downregulation (increased SOCE) decreases Aβ42 accumulation, whereas SOCE inhibition increases Aβ42 accumulation, establishing that ORAI2 negatively regulates SOCE and thereby modulates amyloid-beta production. siRNA knockdown and overexpression, Ca2+ imaging (SOCE measurement), SOCE inhibitor (BTP2), Aβ42 ELISA in H4-APPswe cells International journal of molecular sciences Medium 32722509

Source papers

Stage 0 corpus · 27 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2006 Large store-operated calcium selective currents due to co-expression of Orai1 or Orai2 with the intracellular calcium sensor, Stim1. The Journal of biological chemistry 462 16807233
2017 ORAI2 modulates store-operated calcium entry and T cell-mediated immunity. Nature communications 178 28294127
2015 Upregulated expression of STIM2, TRPC6, and Orai2 contributes to the transition of pulmonary arterial smooth muscle cells from a contractile to proliferative phenotype. American journal of physiology. Cell physiology 104 25673771
2020 ORAI2 Promotes Gastric Cancer Tumorigenicity and Metastasis through PI3K/Akt Signaling and MAPK-Dependent Focal Adhesion Disassembly. Cancer research 95 33310726
2007 Murine ORAI2 splice variants form functional Ca2+ release-activated Ca2+ (CRAC) channels. The Journal of biological chemistry 92 17463004
2023 Peritoneal high-fat environment promotes peritoneal metastasis of gastric cancer cells through activation of NSUN2-mediated ORAI2 m5C modification. Oncogene 61 37130916
2016 Orai1 and Orai2 mediate store-operated calcium entry that regulates HL60 cell migration and FAK phosphorylation. Biochimica et biophysica acta. Molecular cell research 47 27865925
2017 Deletion of Orai2 augments endogenous CRAC currents and degranulation in mast cells leading to enhanced anaphylaxis. Cell calcium 45 29604961
2020 ORAI1 and ORAI2 modulate murine neutrophil calcium signaling, cellular activation, and host defense. Proceedings of the National Academy of Sciences of the United States of America 41 32929002
2015 Orai1-Orai2 complex is involved in store-operated calcium entry in chondrocyte cell lines. Cell calcium 40 25769459
2019 Loss of Orai2-Mediated Capacitative Ca2+ Entry Is Neuroprotective in Acute Ischemic Stroke. Stroke 38 31551038
2020 Orai-1 and Orai-2 regulate oral cancer cell migration and colonisation by suppressing Akt/mTOR/NF-κB signalling. Life sciences 37 32882268
2021 Orai2 Modulates Store-Operated Ca2+ Entry and Cell Cycle Progression in Breast Cancer Cells. Cancers 25 35008277
2015 Regulation of store-operated Ca2+ entry activity by cell cycle dependent up-regulation of Orai2 in brain capillary endothelial cells. Biochemical and biophysical research communications 25 25748572
2020 ORAI2 Down-Regulation Potentiates SOCE and Decreases Aβ42 Accumulation in Human Neuroglioma Cells. International journal of molecular sciences 19 32722509
2014 Orai-2 is localized on secretory granules and regulates antigen-evoked Ca²⁺ mobilization and exocytosis in mast cells. Biochemical and biophysical research communications 15 25044118
2019 Identification of Key Pathways and Genes in the Orai2 Mediated Classical and Mesenchymal Subtype of Glioblastoma by Bioinformatic Analyses. Disease markers 13 31772693
2021 Orai1- and Orai2-, but not Orai3-mediated ICRAC is regulated by intracellular pH. The Journal of physiology 10 34877682
2024 Sodium Houttuyfonate Alleviates Monocrotaline-induced Pulmonary Hypertension by Regulating Orai1 and Orai2. American journal of respiratory cell and molecular biology 9 38709251
2019 Leucine-rich repeat kinase 2 regulates mouse dendritic cell migration by ORAI2. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 9 31166814
2022 Orai2 channel regulates prostaglandin E2 production in TNFα/IL1α-stimulated astrocytes. Glia 8 35506586
2024 ORAI2 is Important for the Development of Early-Stage Postirradiation Fibrosis in Salivary Glands. International journal of radiation oncology, biology, physics 6 39384103
2021 X-Ray Causes mRNA Transcripts Change to Enhance Orai2-Mediated Ca2+ Influx in Rat Brain Microvascular Endothelial Cells. Frontiers in molecular biosciences 5 34595206
2023 Bisbenzylisoquinoline alkaloids from Plumula Nelumbinis inhibit vascular smooth muscle cells migration and proliferation by regulating the ORAI2/Akt pathway. Phytochemistry 4 37119920
2023 Correction: Sanchez-Collado et al. Orai2 Modulates Store-Operated Ca2+ Entry and Cell Cycle Progression in Breast Cancer Cells. Cancers 2021, 14, 114. Cancers 1 36831699
2022 Blockade of Platelet Glycoprotein Ibα Augments Neuroprotection in Orai2-Deficient Mice during Middle Cerebral Artery Occlusion. International journal of molecular sciences 1 36012752
2026 Circ72688 Drives Breast Cancer Invasion and Metastasis via the miR-654-5p/ORAI2 Axis. Oncology research 0 42065066

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