MIOS

gene
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Also known as FLJ20323MIOSea4Yulink

Summary

MIOS (meiosis regulator for oocyte development, HGNC:21905) is a protein-coding gene on chromosome 7p21.3, encoding GATOR2 complex protein MIOS (Q9NXC5). As a component of the GATOR2 complex, functions as an activator of the amino acid-sensing branch of the mTORC1 signaling pathway. It is a selective cancer dependency (DepMap: 46.4% of cell lines).

Predicted to enable zinc ion binding activity. Involved in cellular response to amino acid starvation; positive regulation of TORC1 signaling; and protein-containing complex localization. Located in several cellular components, including cytosol; lysosomal membrane; and nucleoplasm. Part of GATOR2 complex.

Source: NCBI Gene 54468 — RefSeq curated summary.

At a glance

  • GWAS associations: 1
  • Clinical variants (ClinVar): 132 total
  • Cancer dependency (DepMap): dependent in 46.4% of screened cell lines
  • MANE Select transcript: NM_019005

Identifiers

Gene identifiers

FieldValue
HGNC IDHGNC:21905
Approved symbolMIOS
Namemeiosis regulator for oocyte development
Location7p21.3
Locus typegene with protein product
StatusApproved
AliasesFLJ20323, MIO, Sea4, Yulink
Ensembl geneENSG00000164654
Ensembl biotypeprotein_coding
OMIM615359
Entrez54468

Gene structure

Transcript identifiers

Ensembl transcripts: 37 — 34 protein_coding, 2 retained_intron, 1 protein_coding_CDS_not_defined

ENST00000340080, ENST00000405785, ENST00000433056, ENST00000433635, ENST00000445169, ENST00000456533, ENST00000461907, ENST00000479694, ENST00000493227, ENST00000899856, ENST00000899857, ENST00000899858, ENST00000899859, ENST00000899860, ENST00000899861, ENST00000899862, ENST00000899863, ENST00000899864, ENST00000899865, ENST00000899866, ENST00000899867, ENST00000928619, ENST00000963942, ENST00000963943, ENST00000963944, ENST00000963945, ENST00000963946, ENST00000963947, ENST00000963948, ENST00000963949, ENST00000963950, ENST00000963951, ENST00000963952, ENST00000963953, ENST00000963954, ENST00000963955, ENST00000963956

RefSeq mRNA: 5 — MANE Select: NM_019005 NM_001370076, NM_001370077, NM_001370078, NM_001370080, NM_019005

CCDS: CCDS43554

Canonical transcript exons

ENST00000340080 — 13 exons

ExonStartEnd
ENSE0000067125675740987574196
ENSE0000108635275724367573769
ENSE0000127759976069967608932
ENSE0000153814075680267568123
ENSE0000153814275676077567688
ENSE0000182109475668847567072
ENSE0000349651176059427606071
ENSE0000350499275894057589563
ENSE0000352404675856367585805
ENSE0000359481075831187583372
ENSE0000361415675884987588563
ENSE0000365958875962577596461
ENSE0000366187875949807595132

Expression profiles

Bgee: expression breadth ubiquitous, 290 present calls, max score 96.26.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 14.4414 / max 217.0966, expressed in 1788 samples.

FANTOM5 promoters (6 alternative TSS)

Promoter IDTPM avgSamples expressed
771977.65931695
771993.67901480
771982.00111186
772000.8949578
2043490.186037
772010.02123

Top tissues by expression

295 total, by Bgee expression score (0-100, higher = more expressed):

TissueAnatomy IDExpression scoreQuality
tongue squamous epitheliumUBERON:000691996.26gold quality
gastrocnemiusUBERON:000138895.35gold quality
spermCL:000001995.01gold quality
muscle of legUBERON:000138394.63gold quality
pancreatic ductal cellCL:000207994.29gold quality
tibialis anteriorUBERON:000138594.03gold quality
endothelial cellCL:000011593.97gold quality
gingival epitheliumUBERON:000194993.91gold quality
deltoidUBERON:000147693.77gold quality
squamous epitheliumUBERON:000691493.63gold quality
muscle organUBERON:000163093.48gold quality
esophagus squamous epitheliumUBERON:000692093.40gold quality
male germ cellCL:000001593.16gold quality
skeletal muscle tissue of rectus abdominisUBERON:000451193.11gold quality
cervix squamous epitheliumUBERON:000692292.54gold quality
tongueUBERON:000172392.52gold quality
tonsilUBERON:000237292.33gold quality
gingivaUBERON:000182892.21gold quality
superior surface of tongueUBERON:000737191.88gold quality
epithelium of esophagusUBERON:000197691.80gold quality
hindlimb stylopod muscleUBERON:000425291.76gold quality
body of tongueUBERON:001187691.71gold quality
cortical plateUBERON:000534391.27gold quality
skeletal muscle tissueUBERON:000113491.21gold quality
pharyngeal mucosaUBERON:000035591.07gold quality
triceps brachiiUBERON:000150990.86gold quality
body of pancreasUBERON:000115090.83gold quality
gluteal muscleUBERON:000200090.80gold quality
quadriceps femorisUBERON:000137790.33gold quality
vastus lateralisUBERON:000137990.26gold quality

Single-cell (SCXA)

Detected in 1 experiment(s), a significant marker in 1.

ExperimentMarker?Max mean expression
E-ANND-3yes4.38

Regulation

Is transcription factor: no

miRNA regulators (miRDB)

40 targeting MIOS, top 30 by miRDB confidence (max_score; target_count = how many genes the miRNA targets in total — lower means more specific):

miRNAMax scoreAvg scoremiRNA target_count
HSA-MIR-1252-5P100.0069.802774
HSA-MIR-518D-5P100.0067.51979
HSA-MIR-518E-5P100.0067.66954
HSA-MIR-518F-5P100.0067.51979
HSA-MIR-519A-5P100.0067.66954
HSA-MIR-519B-5P100.0067.66954
HSA-MIR-519C-5P100.0067.66954
HSA-MIR-520C-5P100.0067.51979
HSA-MIR-522-5P100.0067.66954
HSA-MIR-523-5P100.0067.66954
HSA-MIR-526A-5P100.0067.51979
HSA-MIR-450099.9972.722367
HSA-MIR-806399.9169.763146
HSA-MIR-130599.9171.433443
HSA-MIR-7162-3P99.8968.161682
HSA-MIR-990299.8969.152250
HSA-MIR-526B-3P99.8874.062587
HSA-MIR-1211999.8768.351653
HSA-MIR-202-3P99.8471.411290
HSA-MIR-469899.8471.414303
HSA-MIR-132399.8369.892471
HSA-MIR-548O-3P99.7469.302228
HSA-MIR-4446-5P99.7269.192544
HSA-MIR-4677-5P99.7070.091940
HSA-MIR-7156-5P99.6468.811369
HSA-MIR-612699.6268.09996
HSA-MIR-4649-3P99.5666.901783
HSA-MIR-427399.4567.931206
HSA-MIR-4796-5P99.3470.06810
HSA-MIR-593-3P99.2267.281327

Functional genomics

DepMap (CRISPR cell-line fitness): dependent in 46.4% of screened cell lines.

Cross-species orthologs

5 orthologs

OrganismSymbolGene ID
danio_reriomiosENSDARG00000102587
mus_musculusMiosENSMUSG00000042447
rattus_norvegicusMiosENSRNOG00000007924
drosophila_melanogastermioFBGN0031399
caenorhabditis_elegansWBGENE00018193

Protein

Protein identifiers

GATOR2 complex protein MIOSQ9NXC5 (reviewed: Q9NXC5)

Alternative names: Missing oocyte meiosis regulator homolog

All UniProt accessions (4): C9JAQ1, C9JTV2, C9JUE6, Q9NXC5

UniProt curated annotations — full annotation on UniProt →

Function. As a component of the GATOR2 complex, functions as an activator of the amino acid-sensing branch of the mTORC1 signaling pathway. The GATOR2 complex indirectly activates mTORC1 through the inhibition of the GATOR1 subcomplex. GATOR2 probably acts as an E3 ubiquitin-protein ligase toward GATOR1. In the presence of abundant amino acids, the GATOR2 complex mediates ubiquitination of the NPRL2 core component of the GATOR1 complex, leading to GATOR1 inactivation. In the absence of amino acids, GATOR2 is inhibited, activating the GATOR1 complex. Within the GATOR2 complex, MIOS is required to prevent autoubiquitination of WDR24, the catalytic subunit of the complex. The GATOR2 complex is required for brain myelination.

Subunit / interactions. Component of the GATOR2 subcomplex, composed of MIOS, SEC13, SEH1L, WDR24 and WDR59. The GATOR2 complex interacts with CASTOR1 and CASTOR2; the interaction is negatively regulated by arginine. CASTOR1 and CASTOR2 convey leucine availability via direct interaction with MIOS. The GATOR2 complex interacts with SESN1, SESN2 and SESN3; the interaction is negatively regulated by amino acids. Interacts with SAR1A and SAR1B; the interaction is direct, disrupted by leucine and mediates the interaction of SAR1A or SAR1B with the GATOR2 complex to negatively regulate the TORC1 signaling upon leucine deprivation.

Subcellular location. Lysosome membrane.

Activity regulation. The GATOR2 complex is negatively regulated by the upstream amino acid sensors CASTOR1 and SESN2, which sequester the GATOR2 complex in absence of amino acids. In the presence of abundant amino acids, GATOR2 is released from CASTOR1 and SESN2 and activated.

Similarity. Belongs to the WD repeat mio family.

Isoforms (2)

UniProt IDNamesCanonical?
Q9NXC5-11yes
Q9NXC5-22

RefSeq proteins (5): NP_001357005, NP_001357006, NP_001357007, NP_001357009, NP_061878* (*=MANE)

Domains & families (InterPro)

IDNameType
IPR001680WD40_rptRepeat
IPR015943WD40/YVTN_repeat-like_dom_sfHomologous_superfamily
IPR031488Zn_ribbon_mioDomain
IPR036322WD40_repeat_dom_sfHomologous_superfamily
IPR037593MIOS/Sea4Family
IPR049092MIOS_a-solDomain

Pfam: PF17034, PF21719, PF21720

UniProt features (29 total): binding site 14, repeat 6, modified residue 2, splice variant 2, mutagenesis site 2, zinc finger region 2, chain 1

Structure

Experimental structures (PDB)

5 structures.

PDBMethodResolution (Å)
9LWFELECTRON MICROSCOPY3.41
9OTIELECTRON MICROSCOPY3.5
9LVKELECTRON MICROSCOPY3.59
7UHYELECTRON MICROSCOPY3.66
9LVJELECTRON MICROSCOPY3.82

Predicted structure (AlphaFold)

ModelpLDDTFraction very-high
AF-Q9NXC5-F183.290.53

Functional residue map

Curated UniProt residues grouped by drug-discovery relevance — catalytic, ligand-binding, modification, and mutation-validated positions. Source: UniProtKB sequence features.

Ligand- & substrate-binding residues (14): 740; 775; 778; 788; 827; 830; 830; 832; 835; 838; 849; 854

Post-translational modifications (2): 759, 766

Mutagenesis-validated functional residues (2):

PositionPhenotype
560impaired assembly of the gator2 complex.
785–788impaired amino-acid-mediated mtorc1 activation.

Function

Pathways and Gene Ontology

Reactome pathways

4 pathways

IDPathway
R-HSA-9639288Amino acids regulate mTORC1
R-HSA-2262752Cellular responses to stress
R-HSA-8953897Cellular responses to stimuli
R-HSA-9711097Cellular response to starvation

MSigDB gene sets: 134 (showing top): BROWNE_HCMV_INFECTION_30MIN_DN, MODULE_97, RODRIGUES_THYROID_CARCINOMA_ANAPLASTIC_UP, GOCC_VACUOLAR_MEMBRANE, GOBP_POSITIVE_REGULATION_OF_TOR_SIGNALING, GRAESSMANN_APOPTOSIS_BY_DOXORUBICIN_DN, GOBP_GLIAL_CELL_DEVELOPMENT, MODULE_182, GOBP_NEUROGENESIS, ATGCAGT_MIR217, YY1_Q6, GGCNKCCATNK_UNKNOWN, GOBP_NEGATIVE_REGULATION_OF_INTRACELLULAR_SIGNAL_TRANSDUCTION, GOBP_NEURAL_PRECURSOR_CELL_PROLIFERATION, GOBP_CELLULAR_COMPONENT_ASSEMBLY_INVOLVED_IN_MORPHOGENESIS

GO Biological Process (8): protein-containing complex localization (GO:0031503), cellular response to nutrient levels (GO:0031669), central nervous system myelin formation (GO:0032289), cellular response to amino acid starvation (GO:0034198), oligodendrocyte differentiation (GO:0048709), oligodendrocyte progenitor proliferation (GO:0070444), negative regulation of TORC1 signaling (GO:1904262), positive regulation of TORC1 signaling (GO:1904263)

GO Molecular Function (3): zinc ion binding (GO:0008270), protein binding (GO:0005515), metal ion binding (GO:0046872)

GO Cellular Component (8): nucleoplasm (GO:0005654), cytoplasm (GO:0005737), lysosomal membrane (GO:0005765), cytosol (GO:0005829), cell junction (GO:0030054), GATOR2 complex (GO:0061700), lysosome (GO:0005764), membrane (GO:0016020)

Reactome top-level categories

Rollup of top-3 pathways:

CategoryPathways
Cellular response to starvation1
Cellular responses to stimuli1
Cellular responses to stress1

GO top-level categories

Rollup of top GO terms by namespace:

CategoryTerms
cellular anatomical structure5
TORC1 signaling2
regulation of TORC1 signaling2
macromolecule localization1
response to nutrient levels1
cellular response to stimulus1
central nervous system myelination1
myelin assembly1
cellular response to starvation1
response to amino acid starvation1
central nervous system development1
glial cell differentiation1
gliogenesis1
neural precursor cell proliferation1
negative regulation of TOR signaling1
positive regulation of TOR signaling1
transition metal ion binding1
binding1
cation binding1
nuclear lumen1
intracellular anatomical structure1
lysosome1
lytic vacuole membrane1
cytoplasm1
protein-containing complex1
Seh1-associated complex1
lytic vacuole1

Protein interactions and networks

STRING

1124 interactions, top by confidence (×1000):

Protein AProtein BPartner UniProtScore
MIOSWDR59Q6PJI9998
MIOSWDR24Q96S15998
MIOSSEH1LQ96EE3997
MIOSSEC13P55735996
MIOSNPRL3Q12980840
MIOSNPRL2Q8WTW4827
MIOSDEPDC5O75140809
MIOSRRAGBQ5VZM2717
MIOSRRAGAQ7L523688
MIOSRRAGCQ9HB90668
MIOSITFG2Q969R8649
MIOSKICS2Q96MD2649
MIOSCASTOR1Q8WTX7640
MIOSKPTNQ9Y664618
MIOSCASTOR2A6NHX0617

IntAct

107 interactions, top by confidence:

ABTypeScore
WDR24MIOSpsi-mi:“MI:0915”(physical association)0.940
MIOSWDR24psi-mi:“MI:0915”(physical association)0.940
WDR24SESN2psi-mi:“MI:0914”(association)0.890
SESN2WDR24psi-mi:“MI:0914”(association)0.890
RBBP5KMT2Dpsi-mi:“MI:0914”(association)0.840
MIOSSEC13psi-mi:“MI:0914”(association)0.790
CASTOR1CASTOR1psi-mi:“MI:0914”(association)0.770
PFDN4PFDN6psi-mi:“MI:0914”(association)0.730
CCT2TXNDC9psi-mi:“MI:0914”(association)0.730
SESN2MIOSpsi-mi:“MI:0915”(physical association)0.710
CASTOR1CASTOR2psi-mi:“MI:0914”(association)0.710
CASTOR1MIOSpsi-mi:“MI:0915”(physical association)0.710
WDR24CASTOR1psi-mi:“MI:0914”(association)0.690
WDR24WDR59psi-mi:“MI:0914”(association)0.680

BioGRID (134): MIOS (Affinity Capture-MS), MIOS (Affinity Capture-MS), MIOS (Affinity Capture-MS), MIOS (Affinity Capture-MS), MIOS (Affinity Capture-MS), MIOS (Affinity Capture-MS), MIOS (Affinity Capture-MS), MIOS (Affinity Capture-MS), MIOS (Affinity Capture-MS), MIOS (Affinity Capture-MS), RRAGA (Affinity Capture-Western), RRAGC (Affinity Capture-Western), MIOS (Affinity Capture-Western), MIOS (Affinity Capture-MS), GATSL3 (Affinity Capture-Western)

ESM2 similar proteins: A0JN62, A2RT67, A4IFB6, E1C3P4, F1QEB7, F1R7R1, O35099, O43264, O54921, O94967, P48553, P51593, Q08CL8, Q08CZ0, Q0VEJ0, Q28G25, Q3TLI0, Q4R350, Q5F3K0, Q5R9R1, Q5RFM4, Q5XH48, Q5XIA4, Q5ZIW2, Q641K1, Q6DE97, Q6NU53, Q6VNB8, Q7TMY8, Q7TSG1, Q7Z6Z7, Q8BH15, Q8CGF6, Q8IWR0, Q8IY22, Q8IZQ1, Q8K1X1, Q8N960, Q8QFR2, Q95LL7

Diamond homologs: A4QNS7, Q5RCA2, Q5U5D4, Q802U2, Q8VE19, Q9NXC5, Q9VQ89, B2VZH2, P38164, Q09866, Q2GVT8

SIGNOR signaling

2 interactions.

AEffectBMechanism
MIOS“form complex”GATOR2binding
MIOS“form complex”“CASTOR1-GATOR2 arginine binding complex”binding

Enriched among interaction partners

Reactome pathways and GO biological processes over-represented among this gene’s 87 IntAct physical interaction partners (hypergeometric vs the genome-wide background, BH-FDR, gene-set size 15–500, ranked by fold). A functional readout of the neighbourhood — distinct from this gene’s own memberships above, and biased toward well-studied / hub proteins, so read it as themes rather than proof.

Reactome pathways:

PathwayPartnersFoldFDR
Amino acids regulate mTORC11034.5×7e-11
Prefoldin mediated transfer of substrate to CCT/TriC534.0×6e-05
Cellular response to starvation617.1×2e-04
SUMOylation of DNA damage response and repair proteins512.6×4e-03
Cilium Assembly59.4×9e-03
Organelle biogenesis and maintenance66.8×9e-03
Cellular responses to stress85.1×9e-03

GO biological processes:

GO termPartnersFoldFDR
negative regulation of TORC1 signaling1040.5×2e-11
cellular response to amino acid starvation831.8×3e-08
positive regulation of TORC1 signaling622.2×5e-05
protein folding67.8×8e-03
protein stabilization75.8×1e-02

Disease & clinical

Clinical variants and AI predictions

ClinVar

132 variants total. Per-class counts are floors (≥ shown; pagination cap):

ClassificationCount (floor)
Pathogenic0
Likely pathogenic0
Uncertain significance105
Likely benign4
Benign1

Top pathogenic / likely-pathogenic (0)

SpliceAI

1783 predictions. Top by Δscore:

VariantEffectΔscore
7:7582747:A:Gdonor_gain1.0000
7:7596251:TTGTA:Tacceptor_loss1.0000
7:7596252:TGTA:Tacceptor_loss1.0000
7:7596253:GTAGG:Gacceptor_loss1.0000
7:7596254:TA:Tacceptor_loss1.0000
7:7605936:TCATA:Tacceptor_loss1.0000
7:7605939:TA:Tacceptor_loss1.0000
7:7605940:A:Gacceptor_loss1.0000
7:7605941:G:GAacceptor_loss1.0000
7:7567068:CTCAG:Cdonor_loss0.9900
7:7567069:TCAG:Tdonor_loss0.9900
7:7567071:AGGT:Adonor_loss0.9900
7:7567073:GTC:Gdonor_loss0.9900
7:7567604:AAG:Aacceptor_gain0.9900
7:7568024:A:AGacceptor_gain0.9900
7:7568025:G:GGacceptor_gain0.9900
7:7568025:GTGCT:Gacceptor_gain0.9900
7:7568098:G:GTdonor_gain0.9900
7:7588597:T:Gdonor_gain0.9900
7:7594973:GTTTT:Gacceptor_loss0.9900
7:7594974:TTTTA:Tacceptor_loss0.9900
7:7594975:TTTA:Tacceptor_loss0.9900
7:7594976:TTAG:Tacceptor_loss0.9900
7:7594977:TA:Tacceptor_loss0.9900
7:7594978:AG:Aacceptor_gain0.9900
7:7594979:GG:Gacceptor_gain0.9900
7:7595133:G:GGdonor_gain0.9900
7:7595370:GT:Gdonor_gain0.9900
7:7596255:A:AGacceptor_gain0.9900
7:7596256:G:GGacceptor_gain0.9900

AlphaMissense

5758 scored. Top likely-pathogenic:

VariantProtein changeam_pathogenicity
7:7572711:C:AA79E1.000
7:7572714:T:AV80D1.000
7:7572716:G:AG81R1.000
7:7572716:G:CG81R1.000
7:7572717:G:AG81E1.000
7:7572728:G:CG85R1.000
7:7572728:G:TG85C1.000
7:7572729:G:AG85D1.000
7:7572729:G:TG85V1.000
7:7572735:T:AV87D1.000
7:7572798:T:CF108S1.000
7:7572816:G:CR114P1.000
7:7572821:T:CC116R1.000
7:7572822:G:AC116Y1.000
7:7572823:T:GC116W1.000
7:7572831:T:AL119H1.000
7:7572836:T:AW121R1.000
7:7572836:T:CW121R1.000
7:7572838:G:CW121C1.000
7:7572838:G:TW121C1.000
7:7572864:C:AA130D1.000
7:7572866:G:CA131P1.000
7:7572869:G:CG132R1.000
7:7572870:G:AG132D1.000
7:7572878:A:GK135E1.000
7:7572879:A:CK135T1.000
7:7572879:A:TK135M1.000
7:7572880:G:CK135N1.000
7:7572880:G:TK135N1.000
7:7572885:G:CR137T1.000

dbSNP variants (sampled 300 via entrez): RS1000000245 (7:7598731 G>A,C), RS1000005435 (7:7598567 C>A,G,T), RS1000100277 (7:7602826 A>G), RS1000250949 (7:7575789 C>G), RS1000300538 (7:7583880 TTAAAA>T), RS1000347055 (7:7581245 G>C,T), RS1000361219 (7:7587370 A>C), RS1000425577 (7:7579264 C>A,T), RS1000438272 (7:7598153 G>T), RS1000562929 (7:7576857 A>G), RS1000708800 (7:7582447 G>A), RS1000736245 (7:7589290 G>A), RS1000739877 (7:7582776 G>T), RS1000784593 (7:7605127 A>G), RS1000914816 (7:7594644 T>C)

Disease associations

OMIM: gene MIM:615359 | disease phenotypes:

GenCC curated gene-disease

Mondo (0):

Orphanet (0):

HPO phenotypes

0 total (0 of 0 shown, HPO-id order):

GWAS associations

1 associations (top):

StudyTraitp-value
GCST009391_1481Metabolite levels7.000000e-06

EFO canonical traits (1, from GWAS)

EFO IDTrait name
EFO:0010414triacylglycerol 52:2 measurement

Drugs & pharmacology

Drug and pharmacology data

Is drug target: no

PharmGKB: 1 entry (VIP=true, CPIC=false)

CTD chemical–gene interactions

24 total (human), top 24 by PubMed support.

ChemicalActions (top 5)PubMed papers
sodium arsenitedecreases expression2
Valproic Acidincreases expression, decreases expression2
GSK-J4decreases expression1
dicrotophosdecreases expression1
methylmercuric chloridedecreases expression1
triphenyl phosphateaffects expression1
cobaltous chloridedecreases expression1
abrineincreases expression1
jinfukangdecreases expression1
Resveratrolaffects cotreatment, increases expression1
Temozolomideincreases expression1
Vorinostatincreases expression1
Demecolcinedecreases expression1
Doxorubicindecreases expression1
Ethyl Methanesulfonatedecreases expression1
Formaldehydedecreases expression1
Hydrogen Peroxideaffects expression1
Methyl Methanesulfonatedecreases expression1
Plant Extractsaffects cotreatment, increases expression1
Quercetindecreases expression1
Seleniumdecreases expression1
Tretinoindecreases expression1
Cadmium Chloridedecreases expression1
Copper Sulfatedecreases expression1

Cellosaurus cell lines

2 cell lines: 2 cancer cell line

First 10 cell lines (id-ordered, not curated):

CellosaurusNameCategorySex
CVCL_B1X3Abcam HeLa MIOS KOCancer cell lineFemale
CVCL_SY40HAP1 MIOS (-)Cancer cell lineMale

Clinical trials (associated diseases)

0 trials via MONDO — disease-level, not drug-specific.

No linked Atlas pages yet — the cross-entity mesh grows as the corpus expands.