PPP4R2

gene
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Summary

PPP4R2 (protein phosphatase 4 regulatory subunit 2, HGNC:18296) is a protein-coding gene on chromosome 3p13, encoding Serine/threonine-protein phosphatase 4 regulatory subunit 2 (Q9NY27). Regulatory subunit of serine/threonine-protein phosphatase 4 (PP4). It is a selective cancer dependency (DepMap: 73.6% of cell lines).

The protein encoded by this gene is a regulatory subunit of the serine/threonine-protein phosphatase 4 complex. In addition to being required for efficient DNA double strand break repair, this complex plays a role in organization of microtubules at centrosomes and processing of spliceosomal snRNPs. Several transcript variants encoding different isoforms have been found for this gene.

Source: NCBI Gene 151987 — RefSeq curated summary.

At a glance

  • GWAS associations: 4
  • Clinical variants (ClinVar): 68 total
  • Cancer dependency (DepMap): dependent in 73.6% of screened cell lines
  • MANE Select transcript: NM_174907

Identifiers

Gene identifiers

FieldValue
HGNC IDHGNC:18296
Approved symbolPPP4R2
Nameprotein phosphatase 4 regulatory subunit 2
Location3p13
Locus typegene with protein product
StatusApproved
Ensembl geneENSG00000163605
Ensembl biotypeprotein_coding
OMIM613822
Entrez151987

Gene structure

Transcript identifiers

Ensembl transcripts: 8 — 6 protein_coding, 1 retained_intron, 1 nonsense_mediated_decay

ENST00000356692, ENST00000460360, ENST00000470976, ENST00000476505, ENST00000482242, ENST00000488810, ENST00000495566, ENST00000710398

RefSeq mRNA: 5 — MANE Select: NM_174907 NM_001318025, NM_001318026, NM_001318027, NM_001318028, NM_174907

CCDS: CCDS2917, CCDS82805

Canonical transcript exons

ENST00000356692 — 9 exons

ExonStartEnd
ENSE000010764707306102373061060
ENSE000010764727306367373063747
ENSE000010764737306485273065141
ENSE000010764747306400373064146
ENSE000018570427299681172997071
ENSE000018957427306539773069198
ENSE000035114527305903773059130
ENSE000035925787299807772998158
ENSE000036115947304718673047356

Expression profiles

Bgee: expression breadth ubiquitous, 256 present calls, max score 97.97.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 23.4440 / max 1234.6092, expressed in 1808 samples.

FANTOM5 promoters (1 alternative TSS)

Promoter IDTPM avgSamples expressed
3733223.44401808

Top tissues by expression

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

TissueAnatomy IDExpression scoreQuality
secondary oocyteCL:000065597.97gold quality
oviduct epitheliumUBERON:000480497.73gold quality
epithelial cell of pancreasCL:000008396.83gold quality
adrenal tissueUBERON:001830396.38gold quality
cardiac muscle of right atriumUBERON:000337996.12gold quality
left ventricle myocardiumUBERON:000656695.99gold quality
bone marrowUBERON:000237195.46gold quality
tibialis anteriorUBERON:000138594.67gold quality
germinal epithelium of ovaryUBERON:000130494.59gold quality
mucosa of sigmoid colonUBERON:000499394.49gold quality
oocyteCL:000002394.43gold quality
bone marrow cellCL:000209294.43gold quality
ileal mucosaUBERON:000033194.38gold quality
colonic epitheliumUBERON:000039794.34gold quality
myocardiumUBERON:000234994.33gold quality
islet of LangerhansUBERON:000000694.30gold quality
placentaUBERON:000198794.29gold quality
colonic mucosaUBERON:000031794.15gold quality
deltoidUBERON:000147694.12gold quality
esophagus squamous epitheliumUBERON:000692094.07gold quality
amniotic fluidUBERON:000017394.02gold quality
mucosa of paranasal sinusUBERON:000503094.01gold quality
cartilage tissueUBERON:000241893.91gold quality
tibiaUBERON:000097993.82gold quality
ventricular zoneUBERON:000305393.82gold quality
endometriumUBERON:000129593.69gold quality
pigmented layer of retinaUBERON:000178293.48gold quality
heart right ventricleUBERON:000208093.05gold quality
cerebellar vermisUBERON:000472093.05gold quality
trabecular bone tissueUBERON:000248393.01gold quality

Single-cell (SCXA)

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

ExperimentMarker?Max mean expression
E-ANND-3yes7.32

Regulation

Is transcription factor: no

miRNA regulators (miRDB)

245 targeting PPP4R2, 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-190A-3P100.0080.355520
HSA-MIR-5011-5P100.0083.465820
HSA-LET-7A-3P100.0074.033932
HSA-LET-7B-3P100.0074.083913
HSA-LET-7F-1-3P100.0074.023928
HSA-MIR-98-3P100.0074.083907
HSA-MIR-3924100.0072.092394
HSA-MIR-3613-3P100.0076.367965
HSA-MIR-5692A100.0074.406850
HSA-MIR-200B-3P100.0073.312693
HSA-MIR-200C-3P100.0073.352685
HSA-MIR-429100.0073.442698
HSA-MIR-513A-5P100.0069.772465
HSA-MIR-8485100.0077.574731
HSA-MIR-656-3P100.0072.152788
HSA-MIR-4668-3P100.0068.742635
HSA-MIR-1277-5P100.0073.955056
HSA-MIR-186-5P99.9970.833707
HSA-MIR-548AW99.9972.573559
HSA-MIR-548C-3P99.9974.017587
HSA-MIR-366299.9973.825684
HSA-MIR-428299.9975.366408
HSA-MIR-513B-5P99.9969.962150
HSA-MIR-33A-5P99.9968.621055
HSA-MIR-33B-5P99.9968.581062
HSA-LET-7F-2-3P99.9870.982588
HSA-MIR-1185-1-3P99.9871.042593
HSA-MIR-1185-2-3P99.9871.042593
HSA-MIR-4789-5P99.9870.762721
HSA-MIR-27A-3P99.9872.132955

Functional genomics

DepMap (CRISPR cell-line fitness): dependent in 73.6% of screened cell lines.

Literature-anchored findings (GeneRIF, showing 4)

  • A three-protein PP4 phosphatase complex in mammalian cells, containing PP4C, PP4R2, and PP4R3beta, specifically dephosphorylates ATR-mediated gamma-H2AX generated during DNA replication. (PMID:18614045)
  • PP4R2, a regulatory subunit of PP4, mediates the DNA damage-dependent association between RPA2 and the PP4C catalytic subunit. (PMID:20154705)
  • PPP4R2 affects the survival of neuronal cells in response to DNA damage. (PMID:22559936)
  • Results from a study on gene expression variability markers in early-stage human embryos shows that PPP4R2 is a putative expression variability marker for the 3-day, 8-cell embryo stage. (PMID:26288249)

Cross-species orthologs

0 orthologs

Protein

Protein identifiers

Serine/threonine-protein phosphatase 4 regulatory subunit 2Q9NY27 (reviewed: Q9NY27)

All UniProt accessions (7): Q9NY27, A0AA34QVI2, C9IZ04, C9IZF4, F8WCA1, F8WDK3, H7C559

UniProt curated annotations — full annotation on UniProt →

Function. Regulatory subunit of serine/threonine-protein phosphatase 4 (PP4). May regulate the activity of PPP4C at centrosomal microtubule organizing centers. Its interaction with the SMN complex leads to enhance the temporal localization of snRNPs, suggesting a role of PPP4C in maturation of spliceosomal snRNPs. The PPP4C-PPP4R2-PPP4R3A PP4 complex specifically dephosphorylates H2AX phosphorylated on ‘Ser-140’ (gamma-H2AX) generated during DNA replication and required for DNA double strand break repair. Mediates RPA2 dephosphorylation by recruiting PPP4C to RPA2 in a DNA damage-dependent manner. RPA2 dephosphorylation is required for the efficient RPA2-mediated recruitment of RAD51 to chromatin following double strand breaks, an essential step for DNA repair.

Subunit / interactions. Serine/threonine-protein phosphatase 4 (PP4) occurs in different assemblies of the catalytic and one or more regulatory subunits. Component of the PP4 complexes PPP4C-PPP4R2, PPP4C-PPP4R2-PPP4R3A and PPP4C-PPP4R2-PPP4R3B. The PPP4C-PPP4R2 complex appears to be a tetramer composed of 2 molecules of PPP4C and 2 molecules of PPP4R2. Interacts with DDX20/GEMIN3 and GEMIN4. Interacts with RPA2; this DNA damage-dependent interaction recruits PPP4C leading to RPA2 dephosphorylation.

Subcellular location. Cytoplasm. Cytoskeleton. Microtubule organizing center. Centrosome. Nucleus.

Tissue specificity. Widely expressed.

Similarity. Belongs to the PPP4R2 family.

Isoforms (3)

UniProt IDNamesCanonical?
Q9NY27-11yes
Q9NY27-22
Q9NY27-33

RefSeq proteins (5): NP_001304954, NP_001304955, NP_001304956, NP_001304957, NP_777567* (*=MANE)

Domains & families (InterPro)

IDNameType
IPR015267PPP4R2Family

Pfam: PF09184

UniProt features (26 total): compositionally biased region 15, mutagenesis site 3, modified residue 2, splice variant 2, sequence variant 2, chain 1, region of interest 1

Structure

Experimental structures (PDB)

0 structures.

Predicted structure (AlphaFold)

ModelpLDDTFraction very-high
AF-Q9NY27-F162.820.27

Functional residue map

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

Post-translational modifications (2): 159, 226

Mutagenesis-validated functional residues (3):

PositionPhenotype
99no effect on rpa2-binding; decrease in ppp4c-binding.
103no effect on rpa2-binding; loss of ppp4c-binding.
106no effect on rpa2-binding, nor on ppp4c-binding.

Function

Pathways and Gene Ontology

Reactome pathways

1 pathways

IDPathway
R-HSA-5693607Processing of DNA double-strand break ends

MSigDB gene sets: 245 (showing top): GSE45365_NK_CELL_VS_CD8_TCELL_UP, GOBP_REGULATION_OF_DOUBLE_STRAND_BREAK_REPAIR, GOBP_REGULATION_OF_DNA_RECOMBINATION, MODULE_255, MODULE_45, MODULE_317, GOBP_REGULATION_OF_DOUBLE_STRAND_BREAK_REPAIR_VIA_HOMOLOGOUS_RECOMBINATION, GOBP_REGULATION_OF_DNA_REPAIR, GOCC_MICROTUBULE_ORGANIZING_CENTER, MODULE_66, MODULE_118, MODULE_379, chr3p13, MILI_PSEUDOPODIA_HAPTOTAXIS_UP, KMCATNNWGGA_UNKNOWN

GO Biological Process (5): mRNA processing (GO:0006397), RNA splicing (GO:0008380), regulation of double-strand break repair via homologous recombination (GO:0010569), protein modification process (GO:0036211), regulation of double-strand break repair (GO:2000779)

GO Molecular Function (3): protein phosphatase regulator activity (GO:0019888), protein-macromolecule adaptor activity (GO:0030674), protein binding (GO:0005515)

GO Cellular Component (7): chromatin (GO:0000785), nucleus (GO:0005634), nucleoplasm (GO:0005654), cytoplasm (GO:0005737), centrosome (GO:0005813), protein phosphatase 4 complex (GO:0030289), cytoskeleton (GO:0005856)

Reactome top-level categories

Rollup of top-1 pathways:

CategoryPathways
HDR through Homologous Recombination (HRR) or Single Strand Annealing (SSA)1

GO top-level categories

Rollup of top GO terms by namespace:

CategoryTerms
cellular anatomical structure3
RNA processing2
mRNA metabolic process1
regulation of DNA recombination1
double-strand break repair via homologous recombination1
regulation of double-strand break repair1
protein metabolic process1
macromolecule modification1
regulation of DNA repair1
double-strand break repair1
phosphoprotein phosphatase activity1
phosphatase regulator activity1
protein phosphatase binding1
protein binding1
molecular adaptor activity1
binding1
chromosome1
intracellular membrane-bounded organelle1
nuclear lumen1
intracellular anatomical structure1
centriole1
microtubule organizing center1
protein serine/threonine phosphatase complex1
intracellular membraneless organelle1

Protein interactions and networks

STRING

1890 interactions, top by confidence (×1000):

Protein AProtein BPartner UniProtScore
PPP4R2PPP4CP33172998
PPP4R2PPP4R3AQ6IN85995
PPP4R2PPP4R1Q8TF05971
PPP4R2PPP4R3BQ5MIZ7954
PPP4R2PPP4R4Q6NUP7774
PPP4R2GEMIN4P57678725
PPP4R2DDX20Q9UHI6714
PPP4R2PPP6CO00743646
PPP4R2PPP6R2O75170599
PPP4R2PPP6R1Q9UPN7519
PPP4R2PPP4R3CQ6ZMV5498
PPP4R2MAP2K1Q02750455
PPP4R2PPP2CAP05323446
PPP4R2RPA2P15927443
PPP4R2GXYLT2A0PJZ3441

IntAct

82 interactions, top by confidence:

ABTypeScore
PPP4R2PPP4Cpsi-mi:“MI:0914”(association)0.950
PPP4R2PPP4Cpsi-mi:“MI:0915”(physical association)0.950
PPP4CPPP4R2psi-mi:“MI:0915”(physical association)0.950
PPME1PPP2R1Apsi-mi:“MI:0914”(association)0.950
PPP4R3APPP4Cpsi-mi:“MI:0914”(association)0.920
PPP4CPPP4R3Apsi-mi:“MI:0914”(association)0.920
PPP4R3BPPP4Cpsi-mi:“MI:0914”(association)0.890
PPME1PPP2CApsi-mi:“MI:0914”(association)0.880
TIPRLPPP4Cpsi-mi:“MI:0914”(association)0.850
PPP4R2TIPRLpsi-mi:“MI:0914”(association)0.800
PPP4CTCP1psi-mi:“MI:0914”(association)0.730
SNRPD2GEMIN2psi-mi:“MI:0914”(association)0.710
PPP4CSUPT5Hpsi-mi:“MI:0914”(association)0.640
DNAJC8SF3B1psi-mi:“MI:0914”(association)0.640
SF3B1SAP18psi-mi:“MI:0914”(association)0.640
SNRPA1U2SURPpsi-mi:“MI:0914”(association)0.640
DHX38DHX16psi-mi:“MI:0914”(association)0.630
SF3B4SF3B1psi-mi:“MI:0914”(association)0.610
USP10ANKRD28psi-mi:“MI:0914”(association)0.610
PPP4R2SF3B1psi-mi:“MI:0914”(association)0.570
PPP4R2SF3B1psi-mi:“MI:2364”(proximity)0.570
MARCKSL1NMT2psi-mi:“MI:0914”(association)0.530

BioGRID (216): PPP4R2 (Affinity Capture-MS), PPP4R2 (Affinity Capture-MS), PPP4R2 (Affinity Capture-MS), PPP4R2 (Affinity Capture-MS), PPP4R2 (Affinity Capture-MS), CUTA (Co-fractionation), DNAJA1 (Co-fractionation), PPP4C (Co-fractionation), PPP4R2 (Co-fractionation), SMEK1 (Co-fractionation), SMEK2 (Co-fractionation), PPP4C (Affinity Capture-MS), SLC4A1AP (Affinity Capture-MS), SMEK1 (Affinity Capture-MS), SMEK2 (Affinity Capture-MS)

ESM2 similar proteins: A0FKI7, A1L1M4, A5X7A0, A7XYJ6, A8MW92, E1BE02, F8VPQ2, O18805, O35613, O94885, P59598, P59808, Q0VGB7, Q13029, Q28IG6, Q4V9H5, Q5M7D6, Q5R9U6, Q5RJ80, Q5TJE1, Q5XG73, Q5ZHQ6, Q5ZM88, Q5ZMU6, Q63755, Q640U0, Q641E3, Q6DCQ0, Q6I6G8, Q6KAQ7, Q6P964, Q6ZQ03, Q76L83, Q76LL6, Q76N89, Q7TT18, Q8BLG0, Q8BZ32, Q8C9B9, Q8CCJ9

Diamond homologs: A1L1M4, Q0VGB7, Q28IG6, Q5M7D6, Q5R9U6, Q5ZMU6, Q6DCQ0, Q6P964, Q9NY27, Q9W2U4, P0C5B7

SIGNOR signaling

0 interactions.

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
mRNA Splicing - Minor Pathway931.0×6e-10
mRNA Splicing1423.6×7e-14
mRNA Polyadenylation1418.9×1e-12
CHD1 and CHD2 subfamily1118.4×8e-10
Processing of Capped Intron-Containing Pre-mRNA1417.7×2e-12
mRNA Splicing - Major Pathway2016.8×5e-17
Influenza Infection513.5×1e-03
Metabolism of RNA2012.8×5e-15

GO biological processes:

GO termPartnersFoldFDR
U2-type prespliceosome assembly1079.0×5e-15
RNA splicing, via transesterification reactions647.4×2e-07
mRNA splicing, via spliceosome1719.7×4e-15
RNA splicing1213.4×1e-08

Disease & clinical

Clinical variants and AI predictions

ClinVar

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

ClassificationCount (floor)
Pathogenic0
Likely pathogenic0
Uncertain significance48
Likely benign3
Benign0

Top pathogenic / likely-pathogenic (0)

SpliceAI

2177 predictions. Top by Δscore:

VariantEffectΔscore
3:72997059:G:GTdonor_gain1.0000
3:72998071:A:AGacceptor_gain1.0000
3:72998072:TCTA:Tacceptor_loss1.0000
3:72998073:CTA:Cacceptor_loss1.0000
3:72998075:A:AGacceptor_gain1.0000
3:72998075:AGA:Aacceptor_loss1.0000
3:72998076:G:GAacceptor_gain1.0000
3:72998076:GA:Gacceptor_gain1.0000
3:72998076:GAT:Gacceptor_gain1.0000
3:72998076:GATT:Gacceptor_gain1.0000
3:72998076:GATTT:Gacceptor_gain1.0000
3:72998154:ACAAT:Adonor_gain1.0000
3:72998155:CAATG:Cdonor_loss1.0000
3:72998156:AATGT:Adonor_loss1.0000
3:72998157:AT:Adonor_gain1.0000
3:72998157:ATG:Adonor_loss1.0000
3:72998158:TGT:Tdonor_loss1.0000
3:72998159:G:GGdonor_gain1.0000
3:72998159:GTGAG:Gdonor_loss1.0000
3:72998160:T:Gdonor_loss1.0000
3:72998161:GAGTT:Gdonor_loss1.0000
3:73047181:TATA:Tacceptor_loss1.0000
3:73047183:TAG:Tacceptor_loss1.0000
3:73047184:A:AGacceptor_gain1.0000
3:73047184:A:Cacceptor_loss1.0000
3:73047185:G:GGacceptor_gain1.0000
3:73047322:G:GTdonor_gain1.0000
3:73047323:A:Tdonor_gain1.0000
3:73047355:GG:Gdonor_gain1.0000
3:73047355:GGGT:Gdonor_loss1.0000

AlphaMissense

2789 scored. Top likely-pathogenic:

VariantProtein changeam_pathogenicity
3:72998125:T:CF28S1.000
3:72998128:T:CL29P1.000
3:72998139:G:CA33P1.000
3:72998140:C:AA33D1.000
3:72998148:G:AG36R1.000
3:72998148:G:CG36R1.000
3:72998149:G:AG36E1.000
3:72998149:G:TG36V1.000
3:73047193:T:AW42R1.000
3:73047193:T:CW42R1.000
3:73047194:G:CW42S1.000
3:73047195:G:CW42C1.000
3:73047195:G:TW42C1.000
3:73047215:T:CF49S1.000
3:73047225:A:CK52N1.000
3:73047225:A:TK52N1.000
3:73047227:T:CL53P1.000
3:73047247:T:CF60L1.000
3:73047248:T:CF60S1.000
3:73047248:T:GF60C1.000
3:73047249:C:AF60L1.000
3:73047249:C:GF60L1.000
3:73047293:T:AV75D1.000
3:73047307:T:CF80L1.000
3:73047309:T:AF80L1.000
3:73047309:T:GF80L1.000
3:73047321:G:CK84N1.000
3:73047321:G:TK84N1.000
3:73047329:T:AI87K1.000
3:73047329:T:GI87R1.000

dbSNP variants (sampled 300 via entrez): RS1000001682 (3:73033817 C>T), RS1000100460 (3:73038851 T>C), RS1000104804 (3:73025158 A>G), RS1000139143 (3:73044443 G>A,C), RS1000176054 (3:73005872 A>G,T), RS1000196140 (3:73040058 G>GA), RS1000212310 (3:73012593 G>A), RS1000232733 (3:73010554 C>G,T), RS1000246206 (3:73044185 A>T), RS1000251805 (3:72996914 C>A,G,T), RS1000253956 (3:73044269 C>T), RS1000267381 (3:73010348 C>T), RS1000279281 (3:73062983 G>A,C), RS1000306728 (3:73010089 T>C), RS1000397764 (3:73048977 C>G,T)

Disease associations

OMIM: gene MIM:613822 | disease phenotypes:

GenCC curated gene-disease

Mondo (0):

Orphanet (0):

HPO phenotypes

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

GWAS associations

4 associations (top):

StudyTraitp-value
GCST006626_45Pulse pressure3.000000e-10
GCST007269_29Pulse pressure7.000000e-09
GCST007392_2Mitochondrial DNA copy number (white blood cells)6.000000e-07
GCST010002_431Refractive error3.000000e-11

EFO canonical traits (2, from GWAS)

EFO IDTrait name
EFO:0005763pulse pressure measurement
EFO:0006312mitochondrial DNA measurement

Drugs & pharmacology

Drug and pharmacology data

Is drug target: no

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

CTD chemical–gene interactions

54 total (human), top 30 by PubMed support.

ChemicalActions (top 5)PubMed papers
Valproic Acidaffects cotreatment, decreases expression, decreases methylation7
Cadmiumdecreases reaction, increases abundance, increases expression2
Formaldehydedecreases expression2
Phenylmercuric Acetateaffects cotreatment, decreases expression2
Tobacco Smoke Pollutionincreases expression2
aristolochic acid Idecreases expression1
FR900359affects phosphorylation1
dicrotophosdecreases expression1
2,4,6-tribromophenoldecreases expression1
methylmercuric chloridedecreases expression1
triphenyl phosphateaffects expression1
bisphenol Aaffects expression1
decabromobiphenyl etherincreases expression1
beta-lapachonedecreases expression1
arseniteaffects binding, decreases reaction1
tetrabromobisphenol Adecreases expression1
benzo(e)pyrenedecreases methylation1
coumarinaffects phosphorylation1
K 7174increases expression1
4-(5-benzo(1,3)dioxol-5-yl-4-pyridin-2-yl-1H-imidazol-2-yl)benzamideaffects cotreatment, decreases expression1
2,2’,4,4’-tetrabromodiphenyl etherincreases expression1
dorsomorphinaffects cotreatment, decreases expression1
Resveratrolincreases expression, affects cotreatment1
Sunitinibincreases expression1
Leflunomideincreases expression1
Cidofovirdecreases expression1
Acetaminophendecreases expression1
Air Pollutantsdecreases expression, increases abundance1
Azathioprineincreases expression1
Benzo(a)pyrenedecreases expression1

Cellosaurus cell lines

3 cell lines: 2 cancer cell line, 1 transformed cell line

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

CellosaurusNameCategorySex
CVCL_B3ELAbcam HEK293T PPP4R2 KOTransformed cell lineFemale
CVCL_TG24HAP1 PPP4R2 (-) 1Cancer cell lineMale
CVCL_TG25HAP1 PPP4R2 (-) 2Cancer 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.