PITPNA

gene
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Also known as VIB1A

Summary

PITPNA (phosphatidylinositol transfer protein alpha, HGNC:9001) is a protein-coding gene on chromosome 17p13.3, encoding Phosphatidylinositol transfer protein alpha isoform (Q00169). Catalyzes the transfer of phosphatidylinositol (PI) and phosphatidylcholine (PC) between membranes.

This gene encodes a member of a family of lipid-binding proteins that transfer molecules of phosphatidylinositol or phosphatidylcholine between membrane surfaces. The protein is implicated in phospholipase C signaling and in the production of phosphatidylinositol 3,4,5-trisphosphate (PIP3) by phosphoinositide-3-kinase.

Source: NCBI Gene 5306 — RefSeq curated summary.

At a glance

  • GWAS associations: 2
  • Clinical variants (ClinVar): 38 total
  • Dosage sensitivity (ClinGen): haploinsufficiency no evidence, triplosensitivity no evidence
  • MANE Select transcript: NM_006224

Identifiers

Gene identifiers

FieldValue
HGNC IDHGNC:9001
Approved symbolPITPNA
Namephosphatidylinositol transfer protein alpha
Location17p13.3
Locus typegene with protein product
StatusApproved
AliasesVIB1A
Ensembl geneENSG00000174238
Ensembl biotypeprotein_coding
OMIM600174
Entrez5306

Gene structure

Transcript identifiers

Ensembl transcripts: 10 — 8 protein_coding, 1 retained_intron, 1 nonsense_mediated_decay

ENST00000313486, ENST00000572004, ENST00000573056, ENST00000573231, ENST00000574991, ENST00000575288, ENST00000575895, ENST00000576010, ENST00000576722, ENST00000576761

RefSeq mRNA: 1 — MANE Select: NM_006224 NM_006224

CCDS: CCDS45563

Canonical transcript exons

ENST00000313486 — 12 exons

ExonStartEnd
ENSE0000119886815625411562792
ENSE0000267133015177181520538
ENSE0000348020815482961548387
ENSE0000349493915585291558559
ENSE0000351683615354411535518
ENSE0000355148315388691538952
ENSE0000357527215215791521645
ENSE0000364053115340991534221
ENSE0000365292115415661541640
ENSE0000366939115530041553149
ENSE0000378727915351821535292
ENSE0000378937215430201543027

Expression profiles

Bgee: expression breadth ubiquitous, 299 present calls, max score 99.69.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 50.3918 / max 431.3695, expressed in 1821 samples.

FANTOM5 promoters (5 alternative TSS)

Promoter IDTPM avgSamples expressed
16373550.35191821
1637330.01613
1637340.01223
1637310.00742
1637320.00411

Top tissues by expression

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

TissueAnatomy IDExpression scoreQuality
pigmented layer of retinaUBERON:000178299.69gold quality
retinaUBERON:000096699.66gold quality
parotid glandUBERON:000183197.55gold quality
ponsUBERON:000098897.28gold quality
skin of legUBERON:000151196.51gold quality
lateral nuclear group of thalamusUBERON:000273696.51gold quality
skin of abdomenUBERON:000141696.26gold quality
monocyteCL:000057696.21gold quality
saliva-secreting glandUBERON:000104496.19gold quality
minor salivary glandUBERON:000183095.98gold quality
prefrontal cortexUBERON:000045195.91gold quality
esophagus mucosaUBERON:000246995.90gold quality
mononuclear cellCL:000084295.89gold quality
mouth mucosaUBERON:000372995.88gold quality
right frontal lobeUBERON:000281095.84gold quality
leukocyteCL:000073895.78gold quality
zone of skinUBERON:000001495.75gold quality
stromal cell of endometriumCL:000225595.69gold quality
gingivaUBERON:000182895.66gold quality
islet of LangerhansUBERON:000000695.39gold quality
rectumUBERON:000105295.34gold quality
gingival epitheliumUBERON:000194995.34gold quality
small intestine Peyer’s patchUBERON:000345495.25gold quality
smooth muscle tissueUBERON:000113595.19gold quality
endothelial cellCL:000011595.10gold quality
frontal cortexUBERON:000187095.10gold quality
frontal lobeUBERON:001652595.10gold quality
dorsolateral prefrontal cortexUBERON:000983495.09gold quality
gall bladderUBERON:000211095.06gold quality
upper arm skinUBERON:000426394.95gold quality

Single-cell (SCXA)

Detected in 4 experiment(s), a significant marker in 3.

ExperimentMarker?Max mean expression
E-GEOD-135922yes15.40
E-CURD-112yes3.83
E-MTAB-8060no138.63
E-ANND-3no0.00

Regulation

Is transcription factor: no

Upstream regulators (CollecTRI, top): TCF3

miRNA regulators (miRDB)

164 targeting PITPNA, 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-4476100.0068.182030
HSA-MIR-6876-5P100.0067.682126
HSA-MIR-3689D100.0066.141181
HSA-MIR-6851-5P100.0065.631294
HSA-MIR-7110-3P100.0073.182486
HSA-MIR-548AW99.9972.573559
HSA-MIR-513B-5P99.9969.962150
HSA-MIR-223-3P99.9970.141140
HSA-MIR-25-3P99.9874.601817
HSA-MIR-32-5P99.9875.211964
HSA-MIR-363-3P99.9874.721821
HSA-MIR-367-3P99.9874.831819
HSA-MIR-92A-3P99.9875.211960
HSA-MIR-92B-3P99.9875.251955
HSA-MIR-103A-3P99.9869.141595
HSA-MIR-10799.9869.141595
HSA-MIR-548AN99.9770.912817
HSA-MIR-1468-3P99.9672.743797
HSA-MIR-211099.9666.681930
HSA-MIR-971899.9468.91918
HSA-MIR-6835-3P99.9370.492904
HSA-MIR-515-5P99.9269.822343
HSA-MIR-519E-5P99.9269.622358
HSA-MIR-497-5P99.9271.832674
HSA-MIR-568099.9169.833421
HSA-MIR-498-3P99.9171.271114
HSA-MIR-106A-5P99.9073.942683
HSA-MIR-15A-5P99.9072.802787
HSA-MIR-15B-5P99.9072.782798
HSA-MIR-16-5P99.9072.802780

Functional genomics

ClinGen dosage: haploinsufficiency 0 (no evidence), triplosensitivity 0 (no evidence). ClinGen Gene Dosage Map

Literature-anchored findings (GeneRIF, showing 6)

  • the PITPalpha structure has to relax to allow access to the Ser166 site, and this may occur at the membrane surface where PI delivery is required for receptor-mediated PLC signaling (PMID:15322105)
  • the apparent occurrence of an unusual TG 3’ splice site in intron 4 is discussed (PMID:17672918)
  • This protein has been found differentially expressed in thalami from patients with schizophrenia. (PMID:20471030)
  • Data show that phosphatidylinositol transfer protein alpha (PITPalpha) is involved in the function of mixed lineage kinase domain-like protein (MLKL) in necroptosis. (PMID:29104146)
  • PDE10A Inhibition Reduces the Manifestation of Pathology in DMD Zebrafish and Represses the Genetic Modifier PITPNA. (PMID:33221436)
  • Restoration of PITPNA in Type 2 diabetic human islets reverses pancreatic beta-cell dysfunction. (PMID:37460527)

Cross-species orthologs

5 orthologs

OrganismSymbolGene ID
danio_reriopitpnaaENSDARG00000039490
danio_reriopitpnabENSDARG00000044091
mus_musculusPitpnaENSMUSG00000017781
rattus_norvegicusPitpnaENSRNOG00000003846
caenorhabditis_elegansWBGENE00021854

Paralogs (5): PITPNM2 (ENSG00000090975), PITPNM3 (ENSG00000091622), PITPNM1 (ENSG00000110697), PITPNC1 (ENSG00000154217), PITPNB (ENSG00000180957)

Protein

Protein identifiers

Phosphatidylinositol transfer protein alpha isoformQ00169 (reviewed: Q00169)

All UniProt accessions (10): Q00169, I3L2X8, I3L3W1, I3L459, I3L471, I3L4C0, I3L4H1, I3L4U7, I3NI10, V9HWC5

UniProt curated annotations — full annotation on UniProt →

Function. Catalyzes the transfer of phosphatidylinositol (PI) and phosphatidylcholine (PC) between membranes. Shows a preference for PI and PC containing shorter saturated or monosaturated acyl chains at the sn-1 and sn-2 positions. Preference order for PC is C16:1 > C16:0 > C18:1 > C18:0 > C20:4 and for PI is C16:1 > C16:0 > C18:1 > C18:0 > C20:4 > C20:3.

Subcellular location. Cytoplasm. Nucleus.

Activity regulation. Phosphatidylinositol transfer activity is inhibited by N-ethylmaleimide.

Similarity. Belongs to the PtdIns transfer protein family. PI transfer class I subfamily.

RefSeq proteins (1): NP_006215* (*=MANE)

Domains & families (InterPro)

IDNameType
IPR001666PI_transferFamily
IPR023393START-like_dom_sfHomologous_superfamily
IPR055261PI_transfer_NDomain

Pfam: PF02121

Catalyzed reactions (Rhea), 2 shown:

  • a 1,2-diacyl-sn-glycero-3-phosphocholine(in) = a 1,2-diacyl-sn-glycero-3-phosphocholine(out) (RHEA:38571)
  • a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol)(in) = a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol)(out) (RHEA:38691)

UniProt features (46 total): helix 12, mutagenesis site 11, strand 8, binding site 6, turn 3, initiator methionine 1, chain 1, modified residue 1, region of interest 1, sequence conflict 1, compositionally biased region 1

Structure

Experimental structures (PDB)

3 structures.

PDBMethodResolution (Å)
9TA5X-RAY DIFFRACTION2.1
8PQOX-RAY DIFFRACTION2.3
1UW5X-RAY DIFFRACTION2.9

Predicted structure (AlphaFold)

ModelpLDDTFraction very-high
AF-Q00169-F195.510.91

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 (6): 58; 60; 85; 89; 96; 194

Post-translational modifications (1): 215

Mutagenesis-validated functional residues (11):

PositionPhenotype
58reduced phosphatidylinositol and phosphatidylcholine transfer activity.
58complete loss of phosphatidylinositol transfer activity but no effect on phosphatidylcholine transfer activity.
58reduced phosphatidylinositol transfer activity but no effect on phosphatidylcholine transfer activity.
60complete loss of phosphatidylinositol transfer activity but no effect on phosphatidylcholine transfer activity.
85reduced phosphatidylinositol transfer activity but no effect on phosphatidylcholine transfer activity.
85reduced phosphatidylinositol and phosphatidylcholine transfer activity.
89significant loss of phosphatidylinositol transfer activity but no effect on phosphatidylcholine transfer activity.
94no effect on phosphatidylinositol transfer activity. resistant to inhibition by n-ethylmaleimide.
102reduced phosphatidylinositol and phosphatidylcholine transfer activity.
187no effect on phosphatidylinositol transfer activity.
202–203significant loss of phosphatidylinositol and phosphatidylcholine transfer activity.

Function

Pathways and Gene Ontology

Reactome pathways

2 pathways

IDPathway
R-HSA-418890Role of second messengers in netrin-1 signaling
R-HSA-8950505Gene and protein expression by JAK-STAT signaling after Interleukin-12 stimulation

MSigDB gene sets: 224 (showing top): TGGTGCT_MIR29A_MIR29B_MIR29C, MORF_MSH3, REACTOME_CYTOKINE_SIGNALING_IN_IMMUNE_SYSTEM, TGCTGCT_MIR15A_MIR16_MIR15B_MIR195_MIR424_MIR497, LFA1_Q6, PID_NETRIN_PATHWAY, GOBP_NEUROGENESIS, SP1_Q2_01, GOBP_ORGANOPHOSPHATE_ESTER_TRANSPORT, GTGCCTT_MIR506, WANG_ESOPHAGUS_CANCER_VS_NORMAL_DN, FONTAINE_PAPILLARY_THYROID_CARCINOMA_UP, GOBP_SENSORY_PERCEPTION_OF_LIGHT_STIMULUS, ATF3_Q6, GOBP_LIPID_METABOLIC_PROCESS

GO Biological Process (6): lipid metabolic process (GO:0006629), axonogenesis (GO:0007409), visual perception (GO:0007601), phospholipid transport (GO:0015914), lipid transport (GO:0006869), intermembrane lipid transfer (GO:0120009)

GO Molecular Function (10): lipid binding (GO:0008289), phosphatidylcholine intramembrane carrier activity (GO:0008525), phosphatidylinositol transfer activity (GO:0008526), phosphatidylcholine binding (GO:0031210), phosphatidylinositol binding (GO:0035091), phosphatidylcholine transfer activity (GO:0120019), phosphatidylglycerol binding (GO:1901611), protein binding (GO:0005515), obsolete phospholipid transporter activity (GO:0005548), quaternary ammonium group binding (GO:0050997)

GO Cellular Component (4): nucleus (GO:0005634), cytoplasm (GO:0005737), cytosol (GO:0005829), extracellular exosome (GO:0070062)

Reactome top-level categories

Rollup of top-2 pathways:

CategoryPathways
Netrin-1 signaling1
Interleukin-12 signaling1

GO top-level categories

Rollup of top GO terms by namespace:

CategoryTerms
lipid transport2
binding2
phospholipid binding2
anion binding2
cellular anatomical structure2
primary metabolic process1
cell morphogenesis involved in neuron differentiation1
neuron projection morphogenesis1
axon development1
sensory perception of light stimulus1
organophosphate ester transport1
transport1
lipid localization1
membrane organization1
intramembrane lipid carrier activity1
phosphatidylinositol binding1
lipid transfer activity1
cation binding1
quaternary ammonium group binding1
phosphatidylcholine binding1
phospholipid transfer activity1
small molecule binding1
intracellular membrane-bounded organelle1
intracellular anatomical structure1
cytoplasm1
extracellular vesicle1

Protein interactions and networks

STRING

1060 interactions, top by confidence (×1000):

Protein AProtein BPartner UniProtScore
PITPNAYWHAEP29360893
PITPNASCARF1Q14162857
PITPNABHLHA9Q7RTU4834
PITPNATRARG1Q8IXB3811
PITPNAPAFAH1B1P43034804
PITPNACRKP46108764
PITPNATIMM22Q9Y584751
PITPNAHIC1Q14526662
PITPNANXF1Q9UBU9591
PITPNASERPINF1P36955560
PITPNAMNTQ99583494
PITPNACDIPTO14735428
PITPNAMYO18AQ92614427
PITPNAOSBPP22059426
PITPNANTN1O95631423
PITPNAPRPH2P23942423

IntAct

40 interactions, top by confidence:

ABTypeScore
PLK1EVI5psi-mi:“MI:0914”(association)0.660
CASP6PITPNApsi-mi:“MI:0915”(physical association)0.560
LAMP2PITPNApsi-mi:“MI:0915”(physical association)0.560
PITPNAUQCRC1psi-mi:“MI:0915”(physical association)0.560
SH3GLB1PITPNApsi-mi:“MI:0915”(physical association)0.560
SIRT3psi-mi:“MI:0915”(physical association)0.400
TK2psi-mi:“MI:0915”(physical association)0.400
CSNK2BPITPNApsi-mi:“MI:0915”(physical association)0.370
Dync1li1SSR3psi-mi:“MI:0914”(association)0.350
Cdc26psi-mi:“MI:0914”(association)0.350
S1PR4PITPNApsi-mi:“MI:0914”(association)0.350
PLEKHG3psi-mi:“MI:0914”(association)0.350
HLA-Cpsi-mi:“MI:0914”(association)0.350
MAPTSHTN1psi-mi:“MI:0914”(association)0.350
LHCGRFURINpsi-mi:“MI:0914”(association)0.350
PITPNBPITPNApsi-mi:“MI:0914”(association)0.350
SSUH2IGLC7psi-mi:“MI:0914”(association)0.350
PITPNBPRKCApsi-mi:“MI:0914”(association)0.350
HSDL1HBBpsi-mi:“MI:0914”(association)0.350
DND1UBA6psi-mi:“MI:0914”(association)0.350

BioGRID (37): PITPNA (Co-fractionation), PITPNA (Co-fractionation), PITPNA (Co-fractionation), PITPNA (Co-fractionation), PITPNA (Co-fractionation), PITPNA (Co-fractionation), PITPNA (Co-fractionation), UBE2L3 (Co-fractionation), PITPNA (Affinity Capture-MS), PITPNA (Affinity Capture-MS), PITPNA (Affinity Capture-MS), PITPNA (Affinity Capture-MS), PMVK (Affinity Capture-MS), MORF4L2 (Affinity Capture-MS), MLKL (Affinity Capture-MS)

ESM2 similar proteins: A8K855, O95299, P0CB89, P0CB90, P11881, P16446, P19824, P29994, P29995, P34942, P48603, P48738, P48739, P53810, P53811, P53812, P60670, P70227, Q00169, Q0E908, Q0MQB6, Q0MQB7, Q14571, Q14573, Q14643, Q14849, Q2HJ54, Q4G005, Q54VC7, Q5BJI9, Q5M7Y0, Q5R6F0, Q63269, Q6IQC7, Q6NRZ4, Q6P3Q6, Q7K556, Q8H1S0, Q8TAT6, Q8WN95

Diamond homologs: G5EEM9, O00562, O35954, O46606, P16446, P43125, P48738, P48739, P53810, P53811, P53812, Q00169, Q28CA0, Q2HJ54, Q3UHE1, Q54D93, Q54VC7, Q5R6F0, Q5U2N3, Q6NZC7, Q6ZPQ6, Q80YA3, Q8K4R4, Q8NEL9, Q8W5R2, Q9BZ71, Q9BZ72, Q9NCL7, Q9NCL8, Q9TR36, Q9U9P7, Q9UKF7, Q12204

SIGNOR signaling

0 interactions.

Disease & clinical

Clinical variants and AI predictions

ClinVar

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

ClassificationCount (floor)
Pathogenic0
Likely pathogenic0
Uncertain significance18
Likely benign1
Benign0

Top pathogenic / likely-pathogenic (0)

SpliceAI

1977 predictions. Top by Δscore:

VariantEffectΔscore
17:1534093:ACTT:Adonor_loss1.0000
17:1534095:TTA:Tdonor_loss1.0000
17:1534096:TACTT:Tdonor_loss1.0000
17:1534097:A:ACdonor_gain1.0000
17:1534097:A:Tdonor_loss1.0000
17:1534097:ACTT:Adonor_gain1.0000
17:1534098:C:CGdonor_gain1.0000
17:1534098:CT:Cdonor_gain1.0000
17:1534098:CTT:Cdonor_gain1.0000
17:1534098:CTTC:Cdonor_gain1.0000
17:1534098:CTTCA:Cdonor_gain1.0000
17:1534100:T:TAdonor_gain1.0000
17:1534218:CTTG:Cacceptor_gain1.0000
17:1534219:TTG:Tacceptor_gain1.0000
17:1534220:TG:Tacceptor_gain1.0000
17:1534222:C:CCacceptor_gain1.0000
17:1534224:A:Cacceptor_gain1.0000
17:1535177:CTTA:Cdonor_loss1.0000
17:1535178:TTAC:Tdonor_loss1.0000
17:1535179:TA:Tdonor_loss1.0000
17:1535180:A:ACdonor_gain1.0000
17:1535180:AC:Adonor_gain1.0000
17:1535180:ACC:Adonor_loss1.0000
17:1535181:C:CAdonor_gain1.0000
17:1535181:CC:Cdonor_gain1.0000
17:1535181:CCT:Cdonor_gain1.0000
17:1535181:CCTT:Cdonor_gain1.0000
17:1535181:CCTTA:Cdonor_gain1.0000
17:1535289:CTTG:Cacceptor_gain1.0000
17:1535298:T:TCacceptor_gain1.0000

AlphaMissense

1797 scored. Top likely-pathogenic:

VariantProtein changeam_pathogenicity
17:1534136:C:GR244P1.000
17:1534164:A:GW235R1.000
17:1534164:A:TW235R1.000
17:1535227:G:CF200L1.000
17:1535227:G:TF200L1.000
17:1535229:A:GF200L1.000
17:1535245:T:AK194N1.000
17:1535245:T:GK194N1.000
17:1541595:G:CH115D1.000
17:1548303:G:CC94W1.000
17:1548318:A:CN89K1.000
17:1548318:A:TN89K1.000
17:1548323:A:GW88R1.000
17:1548323:A:TW88R1.000
17:1548330:C:AE85D1.000
17:1548330:C:GE85D1.000
17:1548370:A:TV72D1.000
17:1553021:C:AK60N1.000
17:1553021:C:GK60N1.000
17:1553023:T:CK60E1.000
17:1553086:C:GG39R1.000
17:1553116:C:GA29P1.000
17:1553133:A:GL23P1.000
17:1553133:A:TL23Q1.000
17:1553139:C:TG21E1.000
17:1534106:A:GL254P0.999
17:1534126:T:AE247D0.999
17:1534126:T:GE247D0.999
17:1534127:T:AE247V0.999
17:1534139:A:TI243N0.999

dbSNP variants (sampled 300 via entrez): RS1000015190 (17:1528241 A>G), RS1000032744 (17:1534763 T>C), RS1000101300 (17:1535769 G>A), RS1000104661 (17:1524334 G>A,T), RS1000195913 (17:1547666 T>A,C), RS1000226387 (17:1544613 G>A), RS1000267913 (17:1560113 G>A), RS1000478608 (17:1544736 G>A,C), RS1000487794 (17:1534352 G>A,T), RS1000515763 (17:1538103 A>G), RS1000604571 (17:1563484 T>A), RS1000651828 (17:1537838 T>A), RS1000677254 (17:1543825 A>T), RS1000797504 (17:1549366 G>T), RS1000815201 (17:1534046 T>C)

Disease associations

OMIM: gene MIM:600174 | disease phenotypes:

GenCC curated gene-disease

Mondo (0):

Orphanet (0):

HPO phenotypes

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

GWAS associations

2 associations (top):

StudyTraitp-value
GCST003830_2Response to bronchodilator in chronic obstructive pulmonary disease (change in FEV1)5.000000e-09
GCST009391_104Metabolite levels7.000000e-06

EFO canonical traits (2, from GWAS)

EFO IDTrait name
EFO:0005921FEV change measurement
EFO:0009767glycine measurement

Drugs & pharmacology

Drug and pharmacology data

Is drug target: no

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

CTD chemical–gene interactions

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

ChemicalActions (top 5)PubMed papers
Valproic Acidaffects cotreatment, increases expression5
bisphenol Adecreases expression, increases expression2
sodium arsenitedecreases expression, increases expression2
Acetaminophenincreases expression2
Rotenonedecreases expression, increases expression2
Tobacco Smoke Pollutionaffects expression, increases expression2
dicrotophosincreases expression1
testosterone enanthateaffects expression1
triphenyl phosphateaffects expression1
propylparabenincreases expression1
trichostatin Aaffects expression1
methylparabenincreases expression1
tris(1,3-dichloro-2-propyl)phosphatedecreases expression1
di-n-butylphosphoric acidaffects expression1
azoxystrobindecreases expression1
CGP 52608affects binding, increases reaction1
deguelindecreases expression1
fenpyroximatedecreases expression1
4-chloro-N-((4-(1,1-dimethylethyl)phenyl)methyl)-3-ethyl-1-methyl-1H-pyrazole-5-carboxamidedecreases expression1
4-(5-benzo(1,3)dioxol-5-yl-4-pyridin-2-yl-1H-imidazol-2-yl)benzamideaffects cotreatment, increases expression1
pyrimidifendecreases expression1
bisphenol Bincreases expression1
pyrachlostrobindecreases expression1
dorsomorphinaffects cotreatment, increases expression1
picoxystrobindecreases expression1
bisphenol AFincreases expression1
Norethindrone Acetateaffects cotreatment, increases expression1
Antimycin Adecreases expression1
Benzo(a)pyreneaffects methylation1
Doxorubicindecreases expression1

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.