INTS4

gene
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Also known as INT4MGC16733MST093

Summary

INTS4 (integrator complex subunit 4, HGNC:25048) is a protein-coding gene on chromosome 11q14.1, encoding Integrator complex subunit 4 (Q96HW7). Component of the integrator complex, a multiprotein complex that terminates RNA polymerase II (Pol II) transcription in the promoter-proximal region of genes. It is a common-essential gene (DepMap: required in 99.2% of cancer cell lines).

INTS4 is a subunit of the Integrator complex, which associates with the C-terminal domain of RNA polymerase II large subunit (POLR2A; MIM 180660) and mediates 3-prime end processing of small nuclear RNAs U1 (RNU1; MIM 180680) and U2 (RNU2; MIM 180690) (Baillat et al., 2005 [PubMed 16239144]).

Source: NCBI Gene 92105 — RefSeq curated summary.

At a glance

  • GWAS associations: 3
  • Clinical variants (ClinVar): 90 total
  • Cancer dependency (DepMap): dependent in 99.2% of screened cell lines (common-essential)
  • MANE Select transcript: NM_033547

Identifiers

Gene identifiers

FieldValue
HGNC IDHGNC:25048
Approved symbolINTS4
Nameintegrator complex subunit 4
Location11q14.1
Locus typegene with protein product
StatusApproved
AliasesINT4, MGC16733, MST093
Ensembl geneENSG00000149262
Ensembl biotypeprotein_coding
OMIM611348
Entrez92105

Gene structure

Transcript identifiers

Ensembl transcripts: 53 — 24 protein_coding, 18 nonsense_mediated_decay, 10 retained_intron, 1 protein_coding_CDS_not_defined

ENST00000433818, ENST00000524766, ENST00000525931, ENST00000527522, ENST00000529807, ENST00000531059, ENST00000533180, ENST00000534064, ENST00000534408, ENST00000535943, ENST00000695096, ENST00000695097, ENST00000695098, ENST00000695099, ENST00000695100, ENST00000695101, ENST00000695102, ENST00000695103, ENST00000695104, ENST00000695105, ENST00000695303, ENST00000695304, ENST00000695305, ENST00000695306, ENST00000695307, ENST00000695308, ENST00000695309, ENST00000695310, ENST00000695311, ENST00000695312, ENST00000695313, ENST00000695314, ENST00000695315, ENST00000695316, ENST00000695317, ENST00000695318, ENST00000695319, ENST00000695320, ENST00000695321, ENST00000695332, ENST00000695333, ENST00000696971, ENST00000889292, ENST00000889293, ENST00000925639, ENST00000925640, ENST00000925641, ENST00000925642, ENST00000925643, ENST00000925644, ENST00000925645, ENST00000942459, ENST00000942460

RefSeq mRNA: 1 — MANE Select: NM_033547 NM_033547

CCDS: CCDS31644

Canonical transcript exons

ENST00000534064 — 23 exons

ExonStartEnd
ENSE000014696527797899677979102
ENSE000014696547799110877991299
ENSE000034820137795594277956062
ENSE000035052297796034177960391
ENSE000035349007798145977981576
ENSE000035610177790771777907810
ENSE000035627267793865177938825
ENSE000035785397789429077894349
ENSE000035800297790354077903620
ENSE000035900927792475077924892
ENSE000035929387790142177901551
ENSE000036058197792134077921473
ENSE000036367677794118077941251
ENSE000036559907792834277928547
ENSE000036608627791882177918978
ENSE000036646207792235677922471
ENSE000036656137795874677958834
ENSE000036677997796095377961138
ENSE000039633927787872077879127
ENSE000039633937799459077994668
ENSE000039634077788383277883952
ENSE000039634117789168177891840
ENSE000039634167789131977891462

Expression profiles

Bgee: expression breadth ubiquitous, 251 present calls, max score 97.42.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 18.1524 / max 221.5980, expressed in 1797 samples.

FANTOM5 promoters (3 alternative TSS)

Promoter IDTPM avgSamples expressed
12145215.59531779
1214531.2936565
1214541.2636414

Top tissues by expression

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

TissueAnatomy IDExpression scoreQuality
buccal mucosa cellCL:000233697.42gold quality
pancreatic ductal cellCL:000207995.50silver quality
adrenal tissueUBERON:001830393.00gold quality
granulocyteCL:000009491.76gold quality
right uterine tubeUBERON:000130291.39gold quality
tibial nerveUBERON:000132390.97gold quality
ganglionic eminenceUBERON:000402390.47gold quality
body of pancreasUBERON:000115090.44gold quality
ventricular zoneUBERON:000305390.33gold quality
spleenUBERON:000210690.26gold quality
left ovaryUBERON:000211990.25gold quality
cerebellar hemisphereUBERON:000224590.13gold quality
right ovaryUBERON:000211890.10gold quality
right hemisphere of cerebellumUBERON:001489090.08gold quality
body of uterusUBERON:000985390.03gold quality
cerebellar cortexUBERON:000212989.92gold quality
bone marrow cellCL:000209289.88gold quality
right lobe of liverUBERON:000111489.80gold quality
endocervixUBERON:000045889.78gold quality
sural nerveUBERON:001548889.75gold quality
ectocervixUBERON:001224989.61gold quality
lymph nodeUBERON:000002989.29gold quality
small intestine Peyer’s patchUBERON:000345489.17gold quality
muscle layer of sigmoid colonUBERON:003580589.10gold quality
right lobe of thyroid glandUBERON:000111989.04gold quality
C1 segment of cervical spinal cordUBERON:000646988.99gold quality
primordial germ cell in gonadCL:0000670 ∩ UBERON:000099188.96gold quality
left lobe of thyroid glandUBERON:000112088.93gold quality
left adrenal gland cortexUBERON:003582588.87gold quality
right adrenal glandUBERON:000123388.83gold quality

Single-cell (SCXA)

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

ExperimentMarker?Max mean expression
E-ANND-3yes4.07

Regulation

Is transcription factor: no

miRNA regulators (miRDB)

9 targeting INTS4, 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-513C-5P99.5068.421730
HSA-MIR-514B-5P99.5068.191766
HSA-MIR-315498.9466.551455
HSA-MIR-313898.4167.53744
HSA-MIR-4768-3P98.1666.022330
HSA-MIR-411-5P97.1166.82601
HSA-MIR-6730-3P97.0367.54889
HSA-MIR-873-3P96.8466.09786
HSA-MIR-55595.9265.25564

Functional genomics

DepMap (CRISPR cell-line fitness): dependent in 99.2% of screened cell lines, common-essential.

Literature-anchored findings (GeneRIF, showing 2)

  • findings demonstrate that the integrator complex is essential for snRNA maturation and Cajal body homeostasis (PMID:22250197)
  • INTS4 is a specific and conserved interaction partner of INTS9/11 that does not interact with either subunit individually. (PMID:29471365)

Cross-species orthologs

5 orthologs

OrganismSymbolGene ID
danio_rerioints4ENSDARG00000102953
mus_musculusInts4ENSMUSG00000025133
rattus_norvegicusInts4ENSRNOG00000012552
drosophila_melanogasterIntS4FBGN0026679
caenorhabditis_elegansints-4WBGENE00012234

Protein

Protein identifiers

Integrator complex subunit 4Q96HW7 (reviewed: Q96HW7)

All UniProt accessions (19): Q96HW7, A0A8C8UVZ7, A0A8Q3SHK8, A0A8Q3SHL8, A0A8Q3SHM4, A0A8Q3SHR8, A0A8Q3SHT4, A0A8Q3SHU3, A0A8Q3SII1, A0A8Q3WKB5, A0A8Q3WKB9, A0A8Q3WKC2, A0A8Q3WKC3, A0A8Q3WKC7, A0A8Q3WKC9, A0A8Q3WKW5, A0A8V8TLR7, E9PIM3, F8WAA7

UniProt curated annotations — full annotation on UniProt →

Function. Component of the integrator complex, a multiprotein complex that terminates RNA polymerase II (Pol II) transcription in the promoter-proximal region of genes. The integrator complex provides a quality checkpoint during transcription elongation by driving premature transcription termination of transcripts that are unfavorably configured for transcriptional elongation: the complex terminates transcription by (1) catalyzing dephosphorylation of the C-terminal domain (CTD) of Pol II subunit POLR2A/RPB1 and SUPT5H/SPT5, (2) degrading the exiting nascent RNA transcript via endonuclease activity and (3) promoting the release of Pol II from bound DNA. The integrator complex is also involved in terminating the synthesis of non-coding Pol II transcripts, such as enhancer RNAs (eRNAs), small nuclear RNAs (snRNAs), telomerase RNAs and long non-coding RNAs (lncRNAs). Within the integrator complex, INTS4 acts as an scaffold that links INTS9 and INTS11. Mediates recruitment of cytoplasmic dynein to the nuclear envelope, probably as component of the integrator complex.

Subunit / interactions. Component of the Integrator complex, composed of core subunits INTS1, INTS2, INTS3, INTS4, INTS5, INTS6, INTS7, INTS8, INTS9/RC74, INTS10, INTS11/CPSF3L, INTS12, INTS13, INTS14 and INTS15. The core complex associates with protein phosphatase 2A subunits PPP2CA and PPP2R1A, to form the Integrator-PP2A (INTAC) complex. INTS4 is part of the RNA endonuclease subcomplex, composed of INTS4, INTS9, INTS11 and inositol hexakisphosphate (InsP6). Interacts with BRAT1; interaction is required for the assembly of the RNA endonuclease subcomplex.

Subcellular location. Nucleus. Cytoplasm.

Miscellaneous. May be produced at very low levels due to a premature stop codon in the mRNA, leading to nonsense-mediated mRNA decay.

Similarity. Belongs to the Integrator subunit 4 family.

Isoforms (3)

UniProt IDNamesCanonical?
Q96HW7-11yes
Q96HW7-22
Q96HW7-33

RefSeq proteins (1): NP_291025* (*=MANE)

Domains & families (InterPro)

IDNameType
IPR011989ARM-likeHomologous_superfamily
IPR016024ARM-type_foldHomologous_superfamily
IPR056235INTS4_8HBDDomain
IPR057412INTS4_CDomain

Pfam: PF13646, PF20168, PF24493, PF25458

UniProt features (92 total): helix 49, strand 20, repeat 8, splice variant 5, turn 3, cross-link 2, mutagenesis site 2, chain 1, sequence conflict 1, modified residue 1

Structure

Experimental structures (PDB)

11 structures.

PDBMethodResolution (Å)
8RC4ELECTRON MICROSCOPY3.1
7CUNELECTRON MICROSCOPY3.5
7BFPELECTRON MICROSCOPY3.56
7PKSELECTRON MICROSCOPY3.6
8RBZELECTRON MICROSCOPY3.7
8R2DELECTRON MICROSCOPY3.9
8RBXELECTRON MICROSCOPY4.1
8YJBELECTRON MICROSCOPY4.1
7BFQELECTRON MICROSCOPY4.15
7YCXELECTRON MICROSCOPY4.18
9VD9ELECTRON MICROSCOPY4.6

Predicted structure (AlphaFold)

ModelpLDDTFraction very-high
AF-Q96HW7-F183.400.49

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 (3): 791, 791, 26

Mutagenesis-validated functional residues (2):

PositionPhenotype
164–167decreased processing activity of the integrator complex.
210decreased processing activity of the integrator complex.

Function

Pathways and Gene Ontology

Reactome pathways

3 pathways

IDPathway
R-HSA-6807505RNA polymerase II transcribes snRNA genes
R-HSA-73857RNA Polymerase II Transcription
R-HSA-74160Gene expression (Transcription)

MSigDB gene sets: 110 (showing top): GOBP_MACROMOLECULE_CATABOLIC_PROCESS, IVANOVA_HEMATOPOIESIS_LATE_PROGENITOR, GOBP_RNA_SURVEILLANCE, GOBP_DNA_TEMPLATED_TRANSCRIPTION_INITIATION, GOBP_TRANSCRIPTION_INITIATION_AT_RNA_POLYMERASE_II_PROMOTER, LASTOWSKA_NEUROBLASTOMA_COPY_NUMBER_DN, NIKOLSKY_BREAST_CANCER_11Q12_Q14_AMPLICON, GOBP_SNRNA_PROCESSING, GOBP_REGULATION_OF_DNA_TEMPLATED_TRANSCRIPTION_ELONGATION, GOCC_INTEGRATOR_COMPLEX, GOCC_NUCLEOLUS, LIU_COMMON_CANCER_GENES, GOBP_DNA_TEMPLATED_TRANSCRIPTION_ELONGATION, TERAO_AOX4_TARGETS_HG_UP, REACTOME_RNA_POLYMERASE_II_TRANSCRIBES_SNRNA_GENES

GO Biological Process (3): snRNA processing (GO:0016180), regulation of transcription elongation by RNA polymerase II (GO:0034243), RNA polymerase II transcription initiation surveillance (GO:0160240)

GO Molecular Function (2): protein-macromolecule adaptor activity (GO:0030674), protein binding (GO:0005515)

GO Cellular Component (6): nucleus (GO:0005634), nucleoplasm (GO:0005654), nucleolus (GO:0005730), cytoplasm (GO:0005737), integrator complex (GO:0032039), INTAC complex (GO:0160232)

Reactome top-level categories

Rollup of top-2 pathways:

CategoryPathways
RNA Polymerase II Transcription1
Gene expression (Transcription)1

GO top-level categories

Rollup of top GO terms by namespace:

CategoryTerms
nuclear lumen2
cellular anatomical structure2
nuclear protein-containing complex2
RNA processing1
snRNA metabolic process1
transcription elongation by RNA polymerase II1
regulation of DNA-templated transcription elongation1
transcription initiation at RNA polymerase II promoter1
nuclear RNA surveillance1
protein binding1
molecular adaptor activity1
binding1
intracellular membrane-bounded organelle1
intracellular membraneless organelle1
intracellular anatomical structure1
integrator complex1

Protein interactions and networks

STRING

1278 interactions, top by confidence (×1000):

Protein AProtein BPartner UniProtScore
INTS4INTS11Q5TA45983
INTS4INTS9Q9NV88937
INTS4KIAA0513O60268915
INTS4INTS3Q68E01890
INTS4INTS1Q8N201888
INTS4INTS5Q6P9B9852
INTS4INTS7Q9NVH2843
INTS4INTS2Q9H0H0831
INTS4INTS12Q96CB8810
INTS4INTS10Q9NVR2807
INTS4WWC1Q8IX03785
INTS4INTS8Q75QN2780
INTS4INTS6Q9UL03744
INTS4INTS13Q9NVM9717
INTS4HAX1O00165679

IntAct

152 interactions, top by confidence:

ABTypeScore
INTS9INTS11psi-mi:“MI:0915”(physical association)0.940
INTS9INTS11psi-mi:“MI:0914”(association)0.940
PPP2R1ASTRNpsi-mi:“MI:0914”(association)0.880
INTS11INTS4psi-mi:“MI:0915”(physical association)0.860
INTS10INTS11psi-mi:“MI:0915”(physical association)0.740
INTS13INTS11psi-mi:“MI:0915”(physical association)0.690
INTS4INTS9psi-mi:“MI:0915”(physical association)0.680
INTS4HGSpsi-mi:“MI:0915”(physical association)0.670
HGSINTS4psi-mi:“MI:0915”(physical association)0.670
FAF2UBBpsi-mi:“MI:0914”(association)0.640
VSIG1TTI1psi-mi:“MI:0914”(association)0.640
PMPCBpsi-mi:“MI:0914”(association)0.640

BioGRID (155): INTS4 (Two-hybrid), INTS4 (Two-hybrid), INTS4 (Two-hybrid), INTS4 (Affinity Capture-MS), INTS4 (Affinity Capture-MS), INTS4 (Affinity Capture-MS), INTS4 (Affinity Capture-MS), INTS4 (Affinity Capture-MS), INTS4 (Affinity Capture-MS), INTS4 (Affinity Capture-MS), INTS4 (Affinity Capture-MS), C7orf26 (Co-fractionation), CPSF3L (Co-fractionation), FAM98A (Co-fractionation), INTS4 (Co-fractionation)

ESM2 similar proteins: A0A1L8F8I9, A0A2R8QPS5, A1A5P5, A7S2N8, B0BM28, B4FGS2, B8AXB6, B8B624, B8JKF4, B9FM64, F1QNV4, F4IQJ2, P49842, P97564, Q08CY4, Q08DB2, Q0P5W1, Q0VA04, Q14AI0, Q2KI89, Q32PH0, Q3SYG4, Q3U0M1, Q4R804, Q5R629, Q61586, Q66I84, Q68F70, Q6DHG8, Q6GL75, Q6GMB0, Q6GN08, Q6GPP1, Q6NU25, Q6PA97, Q7T006, Q7XAM0, Q7Z3E5, Q811G0, Q8CIM8

Diamond homologs: Q54LH5, Q68F70, Q8CIM8, Q96HW7, Q9W3E1

SIGNOR signaling

3 interactions.

AEffectBMechanism
INTS4“up-regulates activity”POLR2Abinding
INTS4“up-regulates quantity”DYNC1H1
INTS4“form complex”“Integrator complex”binding

Enriched among interaction partners

Reactome pathways and GO biological processes over-represented among this gene’s 159 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
FGFR2 mutant receptor activation537.3×7e-06
Spry regulation of FGF signaling535.0×9e-06
Signaling by FGFR2 IIIa TM529.5×2e-05
Abortive elongation of HIV-1 transcript in the absence of Tat629.2×3e-06
Activation of HOX genes during differentiation625.8×7e-06
Pausing and recovery of Tat-mediated HIV elongation725.3×2e-06
Tat-mediated HIV elongation arrest and recovery725.3×2e-06
RNA Pol II CTD phosphorylation and interaction with CE during HIV infection624.0×7e-06

GO biological processes:

GO termPartnersFoldFDR
RNA polymerase II transcription initiation surveillance959.1×7e-12
snRNA processing754.6×5e-09
regulation of transcription elongation by RNA polymerase II847.5×1e-09

Disease & clinical

Clinical variants and AI predictions

ClinVar

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

ClassificationCount (floor)
Pathogenic0
Likely pathogenic0
Uncertain significance70
Likely benign2
Benign0

Top pathogenic / likely-pathogenic (0)

SpliceAI

4410 predictions. Top by Δscore:

VariantEffectΔscore
11:77883827:CTTAC:Cdonor_loss1.0000
11:77883828:TTAC:Tdonor_loss1.0000
11:77883829:TACC:Tdonor_loss1.0000
11:77891329:A:Cdonor_gain1.0000
11:77891675:CCTGA:Cdonor_loss1.0000
11:77891676:CTGA:Cdonor_loss1.0000
11:77891677:TGAC:Tdonor_loss1.0000
11:77891678:GAC:Gdonor_loss1.0000
11:77891679:ACC:Adonor_loss1.0000
11:77901419:A:ACdonor_gain1.0000
11:77901419:AC:Adonor_gain1.0000
11:77901420:C:CCdonor_gain1.0000
11:77901420:CC:Cdonor_gain1.0000
11:77901420:CCCT:Cdonor_gain1.0000
11:77901547:ATAAT:Aacceptor_gain1.0000
11:77901548:TAAT:Tacceptor_gain1.0000
11:77901549:AAT:Aacceptor_gain1.0000
11:77901550:AT:Aacceptor_gain1.0000
11:77901552:C:CAacceptor_loss1.0000
11:77901552:C:CCacceptor_gain1.0000
11:77901553:T:Cacceptor_loss1.0000
11:77901554:A:Cacceptor_gain1.0000
11:77907714:AACCT:Adonor_loss1.0000
11:77907715:A:ATdonor_loss1.0000
11:77907716:C:CTdonor_loss1.0000
11:77918819:A:ATdonor_loss1.0000
11:77918820:C:CAdonor_loss1.0000
11:77918974:GGTAA:Gacceptor_gain1.0000
11:77918975:GTAA:Gacceptor_gain1.0000
11:77918977:AA:Aacceptor_gain1.0000

AlphaMissense

6312 scored. Top likely-pathogenic:

VariantProtein changeam_pathogenicity
11:77883838:A:GW903R1.000
11:77883838:A:TW903R1.000
11:77883870:A:GL892P1.000
11:77891366:C:GA849P1.000
11:77921377:A:GL576P1.000
11:77922454:C:TG511E1.000
11:77922455:C:GG511R1.000
11:77922455:C:TG511R1.000
11:77924855:A:GL470P1.000
11:77924873:C:GR464P1.000
11:77924892:C:GD458H1.000
11:77928418:A:TI432K1.000
11:77928427:A:GL429P1.000
11:77928445:T:AE423V1.000
11:77928448:T:AD422V1.000
11:77928448:T:CD422G1.000
11:77928448:T:GD422A1.000
11:77928449:C:AD422Y1.000
11:77928449:C:GD422H1.000
11:77928450:G:CN421K1.000
11:77928450:G:TN421K1.000
11:77928460:T:GD418A1.000
11:77938662:T:AD385V1.000
11:77938662:T:GD385A1.000
11:77938663:C:GD385H1.000
11:77938665:T:AE384V1.000
11:77938677:A:TV380D1.000
11:77938686:C:TG377E1.000
11:77941201:C:AK323N1.000
11:77941201:C:GK323N1.000

dbSNP variants (sampled 300 via entrez): RS1000011356 (11:77906766 G>T), RS1000072979 (11:77912728 T>C), RS1000105559 (11:77956997 T>A), RS1000147614 (11:77912436 C>CT), RS1000189282 (11:77878553 C>A), RS1000204855 (11:77885736 T>C), RS1000235866 (11:77992402 A>T), RS1000248003 (11:77982583 G>A), RS1000268342 (11:77939859 A>C), RS1000276358 (11:77969441 T>C), RS1000335991 (11:77888485 A>G), RS1000347656 (11:77986538 G>A), RS1000404813 (11:77888161 T>C), RS1000418583 (11:77919830 A>G), RS1000532786 (11:77952103 A>T)

Disease associations

OMIM: gene MIM:611348 | disease phenotypes:

GenCC curated gene-disease

Mondo (0):

Orphanet (0):

HPO phenotypes

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

GWAS associations

3 associations (top):

StudyTraitp-value
GCST007277_15Tourette syndrome5.000000e-06
GCST90002396_472Mean reticulocyte volume2.000000e-09
GCST90002402_382Platelet count2.000000e-11

EFO canonical traits (2, from GWAS)

EFO IDTrait name
EFO:0010701mean reticulocyte volume
EFO:0004309platelet count

Drugs & pharmacology

Drug and pharmacology data

Is drug target: no

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

CTD chemical–gene interactions

23 total (human), top 23 by PubMed support.

ChemicalActions (top 5)PubMed papers
Aflatoxin B1increases methylation2
aristolochic acid Idecreases expression1
TAK-243affects sumoylation1
alpha-pineneaffects cotreatment, increases oxidation, increases abundance1
pirinixic acidaffects binding, increases activity, increases expression1
trichostatin Adecreases expression1
beta-lapachoneincreases expression1
tris(1,3-dichloro-2-propyl)phosphateincreases expression1
methacrylaldehydeincreases oxidation, increases abundance, affects cotreatment1
exemestaneincreases expression1
CGP 52608affects binding, increases reaction1
Acroleinaffects cotreatment, increases oxidation, increases abundance1
Air Pollutantsaffects cotreatment, increases abundance, increases oxidation1
Atrazineincreases expression1
Benzo(a)pyrenedecreases methylation1
Methyl Methanesulfonateincreases expression1
Ozoneincreases abundance, affects cotreatment, increases oxidation1
Dihydrotestosteroneincreases expression1
Valproic Aciddecreases expression1
Cyclosporineincreases expression1
Okadaic Acidincreases expression1
Copper Sulfateincreases expression1
Volatile Organic Compoundsaffects cotreatment, increases oxidation1

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.