KPNA5

gene
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Also known as SRP6IPOA6

Summary

KPNA5 (karyopherin subunit alpha 5, HGNC:6398) is a protein-coding gene on chromosome 6q22.1, encoding Importin subunit alpha-6 (O15131). Functions in nuclear protein import as an adapter protein for nuclear receptor KPNB1.

The transport of molecules between the nucleus and the cytoplasm in eukaryotic cells is mediated by the nuclear pore complex (NPC) which consists of 60-100 proteins and is probably 120 million daltons in molecular size. Small molecules (up to 70 kD) can pass through the nuclear pore by nonselective diffusion; larger molecules are transported by an active process. Most nuclear proteins contain short basic amino acid sequences known as nuclear localization signals (NLSs). KPNA5 protein belongs to the importin alpha protein family and is thought to be involved in NLS-dependent protein import into the nucleus.

Source: NCBI Gene 3841 — RefSeq curated summary.

At a glance

  • GWAS associations: 2
  • Clinical variants (ClinVar): 56 total
  • MANE Select transcript: NM_001366306

Identifiers

Gene identifiers

FieldValue
HGNC IDHGNC:6398
Approved symbolKPNA5
Namekaryopherin subunit alpha 5
Location6q22.1
Locus typegene with protein product
StatusApproved
AliasesSRP6, IPOA6
Ensembl geneENSG00000196911
Ensembl biotypeprotein_coding
OMIM604545
Entrez3841

Gene structure

Transcript identifiers

Ensembl transcripts: 11 — 9 protein_coding, 2 nonsense_mediated_decay

ENST00000356348, ENST00000368564, ENST00000413340, ENST00000697696, ENST00000697697, ENST00000697698, ENST00000697699, ENST00000937497, ENST00000937498, ENST00000967749, ENST00000967750

RefSeq mRNA: 8 — MANE Select: NM_001366306 NM_001366304, NM_001366305, NM_001366306, NM_001366307, NM_001366308, NM_001366309, NM_001366310, NM_002269

CCDS: CCDS5111, CCDS93992, CCDS93993

Canonical transcript exons

ENST00000368564 — 14 exons

ExonStartEnd
ENSE00000762670116702019116702150
ENSE00000762671116705072116705160
ENSE00000762672116716219116716318
ENSE00000762673116722126116722289
ENSE00000762675116725751116725876
ENSE00000762678116729563116729741
ENSE00001888432116726495116726622
ENSE00002360340116724297116724375
ENSE00003609111116689320116689453
ENSE00003971447116698704116698798
ENSE00003971454116692293116692392
ENSE00003971456116732136116741867
ENSE00003971461116692055116692156
ENSE00003971464116681211116681338

Expression profiles

Bgee: expression breadth ubiquitous, 269 present calls, max score 97.41.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 14.8806 / max 401.9012, expressed in 1725 samples.

FANTOM5 promoters (3 alternative TSS)

Promoter IDTPM avgSamples expressed
6943914.39871719
694400.2889152
694380.192977

Top tissues by expression

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

TissueAnatomy IDExpression scoreQuality
calcaneal tendonUBERON:000370197.41gold quality
Brodmann (1909) area 23UBERON:001355493.87gold quality
middle temporal gyrusUBERON:000277193.34gold quality
ponsUBERON:000098892.75gold quality
tibiaUBERON:000097992.60gold quality
bronchial epithelial cellCL:000232892.22gold quality
endothelial cellCL:000011590.91gold quality
pigmented layer of retinaUBERON:000178290.75gold quality
buccal mucosa cellCL:000233690.42gold quality
germinal epithelium of ovaryUBERON:000130490.28gold quality
postcentral gyrusUBERON:000258189.92gold quality
caput epididymisUBERON:000435889.80gold quality
cauda epididymisUBERON:000436089.70gold quality
cortical plateUBERON:000534389.66gold quality
skeletal muscle tissue of biceps brachiiUBERON:000450289.60gold quality
parietal lobeUBERON:000187289.34gold quality
upper leg skinUBERON:000426288.95gold quality
biceps brachiiUBERON:000150788.83gold quality
visceral pleuraUBERON:000240188.63gold quality
superficial temporal arteryUBERON:000161488.45gold quality
mucosa of paranasal sinusUBERON:000503088.33gold quality
superior vestibular nucleusUBERON:000722787.88gold quality
adrenal tissueUBERON:001830387.87gold quality
skin of hipUBERON:000155487.74gold quality
entorhinal cortexUBERON:000272887.64gold quality
corpus epididymisUBERON:000435987.59gold quality
trigeminal ganglionUBERON:000167587.18gold quality
parietal pleuraUBERON:000240087.07gold quality
superior frontal gyrusUBERON:000266186.91gold quality
substantia nigra pars compactaUBERON:000196586.52gold quality

Single-cell (SCXA)

Detected in 9 experiment(s), a significant marker in 4.

ExperimentMarker?Max mean expression
E-CURD-122yes23.11
E-MTAB-9221yes17.78
E-ANND-3yes6.17
E-GEOD-137537yes4.55
E-GEOD-111727no1563.41
E-MTAB-6379no727.59
E-GEOD-83139no3.72
E-MTAB-9801no2.99
E-CURD-112no2.13

Regulation

Is transcription factor: no

miRNA regulators (miRDB)

26 targeting KPNA5, 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-1193100.0065.93529
HSA-MIR-806899.9873.852376
HSA-MIR-433-3P99.9869.371203
HSA-MIR-9983-3P99.9471.483631
HSA-MIR-589-3P99.9169.622088
HSA-MIR-6857-5P99.8765.32985
HSA-MIR-129999.7771.242389
HSA-MIR-4755-5P99.7170.342716
HSA-MIR-5006-3P99.7170.262728
HSA-MIR-3059-5P99.7069.932491
HSA-MIR-6762-3P99.6666.941188
HSA-MIR-150-3P99.4370.51920
HSA-MIR-889-5P99.4168.751025
HSA-MIR-612899.3367.831581
HSA-MIR-412-3P98.8666.89712
HSA-MIR-6754-3P98.8466.60889
HSA-MIR-605-5P98.7968.241161
HSA-MIR-548Q98.7165.35563
HSA-MIR-3928-5P98.5067.48980
HSA-MIR-6806-3P98.5067.31980
HSA-MIR-445098.2668.35725
HSA-MIR-607298.0066.47804
HSA-MIR-134-5P97.1166.52976
HSA-MIR-311897.1166.58984
HSA-MIR-215-3P97.0268.011209
HSA-MIR-6891-3P95.8065.76683

Literature-anchored findings (GeneRIF, showing 4)

  • Study identified the sequences KKKRR, KKKRK, and KKRKK as the optimal sequences for binding to this site for mouse importin-alpha2, human importin-alpha1, and human importin-alpha5, respectively. (PMID:20406804)
  • eVP24 recognizes a unique nonclassical nuclear localization signal binding site on KPNA5 that is necessary for efficient phosphoryated STAT1 nuclear transport. (PMID:25121748)
  • BIG3 may block the KPNAs (KPNA1, KPNA5, and KPNA6) binding region(s) of PHB2. (PMID:26052702)
  • Data suggest that IGFBP5 nuclear import is mediated by KPNA5/KPNB1 complex; nuclear localization sequence of IGFBP5 is critical domain in this nuclear translocation. (IGFBP5 = insulin-like growth factor binding protein-5; KPNA5 = karyopherin subunit alpha-5; KPNB1 = karyopherin subunit beta-1/importin-beta) (PMID:28835592)

Cross-species orthologs

3 orthologs

OrganismSymbolGene ID
danio_reriokpna5ENSDARG00000042030
rattus_norvegicusKpna5ENSRNOG00000030109
drosophila_melanogasterKap-alpha1FBGN0024889

Paralogs (6): KPNA6 (ENSG00000025800), KPNA3 (ENSG00000102753), KPNA1 (ENSG00000114030), KPNA2 (ENSG00000182481), KPNA7 (ENSG00000185467), KPNA4 (ENSG00000186432)

Protein

Protein identifiers

Importin subunit alpha-6O15131 (reviewed: O15131)

Alternative names: Karyopherin subunit alpha-5

All UniProt accessions (6): A0A8V8TL68, A0A8V8TL88, A0A8V8TLL2, A0A8V8TMV2, O15131, Q5TD90

UniProt curated annotations — full annotation on UniProt →

Function. Functions in nuclear protein import as an adapter protein for nuclear receptor KPNB1. Binds specifically and directly to substrates containing either a simple or bipartite NLS motif. Docking of the importin/substrate complex to the nuclear pore complex (NPC) is mediated by KPNB1 through binding to nucleoporin FxFG repeats and the complex is subsequently translocated through the pore by an energy requiring, Ran-dependent mechanism. At the nucleoplasmic side of the NPC, Ran binds to importin-beta and the three components separate and importin-alpha and -beta are re-exported from the nucleus to the cytoplasm where GTP hydrolysis releases Ran from importin. The directionality of nuclear import is thought to be conferred by an asymmetric distribution of the GTP- and GDP-bound forms of Ran between the cytoplasm and nucleus. Mediates nuclear import of STAT1 homodimers and STAT1/STAT2 heterodimers by recognizing non-classical NLSs of STAT1 and STAT2 through ARM repeats 8-9. Recognizes influenza A virus nucleoprotein through ARM repeat 7-9 In vitro, mediates the nuclear import of human cytomegalovirus UL84 by recognizing a non-classical NLS.

Subunit / interactions. Forms a complex with importin subunit beta-1. (Microbial infection) Interacts with ebolavirus protein VP24.

Subcellular location. Cytoplasm.

Tissue specificity. Testis.

Domain organisation. Consists of an N-terminal hydrophilic region, a hydrophobic central region composed of 10 repeats, and a short hydrophilic C-terminus. The N-terminal hydrophilic region contains the importin beta binding domain (IBB domain), which is sufficient for binding importin beta and essential for nuclear protein import. The IBB domain is thought to act as an intrasteric autoregulatory sequence by interacting with the internal autoinhibitory NLS. Binding of KPNB1 probably overlaps the internal NLS and contributes to a high affinity for cytoplasmic NLS-containing cargo substrates. After dissociation of the importin/substrate complex in the nucleus the internal autohibitory NLS contributes to a low affinity for nuclear NLS-containing proteins. The major and minor NLS binding sites are mainly involved in recognition of simple or bipartite NLS motifs. Structurally located within in a helical surface groove they contain several conserved Trp and Asn residues of the corresponding third helices (H3) of ARM repeats which mainly contribute to binding.

Similarity. Belongs to the importin alpha family.

RefSeq proteins (8): NP_001353233, NP_001353234, NP_001353235, NP_001353236, NP_001353237, NP_001353238, NP_001353239, NP_002260 (=MANE)

Domains & families (InterPro)

IDNameType
IPR000225ArmadilloRepeat
IPR002652Importin-a_IBBDomain
IPR011989ARM-likeHomologous_superfamily
IPR016024ARM-type_foldHomologous_superfamily
IPR024931Importin_alphaFamily
IPR032413Arm_3Repeat
IPR036975Importin-a_IBB_sfHomologous_superfamily

Pfam: PF00514, PF01749, PF16186

UniProt features (33 total): helix 12, repeat 10, region of interest 3, sequence variant 2, sequence conflict 2, chain 1, domain 1, short sequence motif 1, compositionally biased region 1

Structure

Experimental structures (PDB)

1 structures.

PDBMethodResolution (Å)
4U2XX-RAY DIFFRACTION3.15

Predicted structure (AlphaFold)

ModelpLDDTFraction very-high
AF-O15131-F185.530.75

Function

Pathways and Gene Ontology

Reactome pathways

11 pathways

IDPathway
R-HSA-1169408ISG15 antiviral mechanism
R-HSA-168276NS1 Mediated Effects on Host Pathways
R-HSA-9918432Maturation of DENV proteins
R-HSA-1169410Antimicrobial mechanism of IFN-stimulated genes
R-HSA-1280215Cytokine Signaling in Immune system
R-HSA-1643685Disease
R-HSA-168255Influenza Infection
R-HSA-168256Immune System
R-HSA-5663205Infectious disease
R-HSA-913531Interferon Signaling
R-HSA-9824446Viral Infection Pathways

MSigDB gene sets: 92 (showing top): WAMUNYOKOLI_OVARIAN_CANCER_LMP_DN, YAGI_AML_WITH_INV_16_TRANSLOCATION, GOBP_INTRACELLULAR_PROTEIN_TRANSPORT, REACTOME_CYTOKINE_SIGNALING_IN_IMMUNE_SYSTEM, BROWNE_HCMV_INFECTION_8HR_UP, BROWNE_HCMV_INFECTION_16HR_UP, GOBP_ESTABLISHMENT_OF_PROTEIN_LOCALIZATION_TO_ORGANELLE, GOBP_NUCLEAR_TRANSPORT, KOYAMA_SEMA3B_TARGETS_UP, DEBIASI_APOPTOSIS_BY_REOVIRUS_INFECTION_UP, DOUGLAS_BMI1_TARGETS_DN, GOBP_PROTEIN_LOCALIZATION_TO_ORGANELLE, GOBP_NLS_BEARING_PROTEIN_IMPORT_INTO_NUCLEUS, GOBP_PROTEIN_LOCALIZATION_TO_NUCLEUS, GOMF_SIGNAL_SEQUENCE_BINDING

GO Biological Process (3): protein import into nucleus (GO:0006606), NLS-bearing protein import into nucleus (GO:0006607), protein transport (GO:0015031)

GO Molecular Function (3): nuclear localization sequence binding (GO:0008139), nuclear import signal receptor activity (GO:0061608), protein binding (GO:0005515)

GO Cellular Component (5): nucleus (GO:0005634), nucleoplasm (GO:0005654), cytosol (GO:0005829), NLS-dependent protein nuclear import complex (GO:0042564), cytoplasm (GO:0005737)

Reactome top-level categories

Rollup of top-9 pathways:

CategoryPathways
Antimicrobial mechanism of IFN-stimulated genes1
Influenza Infection1
Dengue Virus Genome Translation and Replication1
Interferon Signaling1
Immune System1
Viral Infection Pathways1
Disease1
Cytokine Signaling in Immune system1
Infectious disease1

GO top-level categories

Rollup of top GO terms by namespace:

CategoryTerms
cellular anatomical structure3
import into nucleus2
intracellular protein transport1
protein localization to nucleus1
establishment of protein localization to organelle1
protein import into nucleus1
transport1
intracellular protein localization1
establishment of protein localization1
signal sequence receptor activity1
nucleocytoplasmic carrier activity1
binding1
intracellular membrane-bounded organelle1
nuclear lumen1
cytoplasm1
nucleocytoplasmic transport complex1
intracellular anatomical structure1

Protein interactions and networks

STRING

2200 interactions, top by confidence (×1000):

Protein AProtein BPartner UniProtScore
KPNA5KPNB1Q14974887
KPNA5NUP153P49790771
KPNA5STAT1P42224612
KPNA5IPO4Q8TEX9540
KPNA5TNPO1Q92973520
KPNA5RCC1P18754516
KPNA5TNPO2O14787506
KPNA5CSE1LP55060480
KPNA5IPO5O00410475
KPNA5IPO11Q9UI26474
KPNA5IPO7O95373473
KPNA5IPO13O94829470
KPNA5IPO9Q96P70445
KPNA5CAPN2P17655441
KPNA5XPO1O14980437

IntAct

118 interactions, top by confidence:

ABTypeScore
KPNA5NUP50psi-mi:“MI:0915”(physical association)0.830
NUP50KPNA5psi-mi:“MI:0915”(physical association)0.830
NUP50KPNA4psi-mi:“MI:0914”(association)0.830
KPNA5ANP32Bpsi-mi:“MI:0915”(physical association)0.780
ANP32BKPNA5psi-mi:“MI:0915”(physical association)0.780
NUP50KPNA3psi-mi:“MI:0914”(association)0.780
SPOPLKPNA5psi-mi:“MI:0915”(physical association)0.740
KPNA5SPOPLpsi-mi:“MI:0915”(physical association)0.740
CDKN1BKPNA5psi-mi:“MI:0914”(association)0.720
CDKN1BKPNA5psi-mi:“MI:0915”(physical association)0.720
KPNA5CDKN1Bpsi-mi:“MI:0915”(physical association)0.720
CFTRESYT2psi-mi:“MI:0914”(association)0.710
KPNA1TCERG1psi-mi:“MI:0914”(association)0.640
ORF4bKPNA3psi-mi:“MI:0914”(association)0.620
CDKN1BKPNA3psi-mi:“MI:0915”(physical association)0.580
KPNA5POLR3Cpsi-mi:“MI:0915”(physical association)0.560
KPNA5NMNAT1psi-mi:“MI:0915”(physical association)0.560
KPNA5RAD52psi-mi:“MI:0915”(physical association)0.560

BioGRID (116): KPNA5 (Two-hybrid), ANP32B (Two-hybrid), NUP50 (Two-hybrid), SPOPL (Two-hybrid), KPNA5 (Affinity Capture-RNA), KPNA5 (Affinity Capture-RNA), KPNA5 (Affinity Capture-MS), KPNA5 (Affinity Capture-MS), KPNA5 (Affinity Capture-MS), KPNA5 (Co-fractionation), KPNA5 (Co-fractionation), KPNA5 (Co-fractionation), KPNA5 (Co-fractionation), KPNA5 (Two-hybrid), KPNA5 (Affinity Capture-MS)

ESM2 similar proteins: A2VE08, A7Y521, E1C6Q1, O15131, O35116, O35142, O35345, O55029, O60684, O70133, O88544, O94973, P17427, P18484, P26233, P35605, P35606, P52294, P52297, P70188, P83953, Q08211, Q0V7M0, Q0VCK5, Q13098, Q28141, Q3SZA0, Q4R5E6, Q503E9, Q56R16, Q5F418, Q5R648, Q5R664, Q5R874, Q5R909, Q5RBV0, Q5ZHN3, Q5ZML1, Q60960, Q68FK8

Diamond homologs: A2VE08, A9QM74, B6HJ92, C0LLJ0, C1JZ66, C6K7I2, F4JL11, G5EB89, O00505, O00629, O04294, O14063, O15131, O22478, O35343, O35344, O35345, O60684, O80480, O94374, P52170, P52171, P52292, P52293, P52294, P52295, P83953, P91276, Q02821, Q0V7M0, Q19969, Q503E9, Q557F4, Q56R16, Q5R909, Q5RBV0, Q5ZML1, Q60960, Q71VM4, Q76P29

SIGNOR signaling

0 interactions.

Enriched among interaction partners

Reactome pathways and GO biological processes over-represented among this gene’s 59 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
NS1 Mediated Effects on Host Pathways645.1×1e-06
ISG15 antiviral mechanism623.7×3e-05
Metabolism of RNA66.6×6e-03
Cytokine Signaling in Immune system66.4×6e-03
Cell Cycle65.7×9e-03

GO biological processes:

GO termPartnersFoldFDR
protein import into nucleus721.0×2e-05
regulation of apoptotic process813.9×2e-05
RNA splicing712.9×1e-04
mRNA processing69.8×2e-03
proteasome-mediated ubiquitin-dependent protein catabolic process77.6×2e-03

Disease & clinical

Clinical variants and AI predictions

ClinVar

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

ClassificationCount (floor)
Pathogenic0
Likely pathogenic0
Uncertain significance45
Likely benign0
Benign0

Top pathogenic / likely-pathogenic (0)

SpliceAI

2334 predictions. Top by Δscore:

VariantEffectΔscore
6:116681337:GG:Gdonor_gain1.0000
6:116681338:GG:Gdonor_gain1.0000
6:116681339:G:GGdonor_gain1.0000
6:116682992:A:AGdonor_gain1.0000
6:116689315:TTAA:Tacceptor_loss1.0000
6:116689316:TAA:Tacceptor_loss1.0000
6:116689317:A:AGacceptor_gain1.0000
6:116689317:AAGAT:Aacceptor_gain1.0000
6:116689318:A:Cacceptor_loss1.0000
6:116689318:A:Gacceptor_gain1.0000
6:116689318:AGAT:Aacceptor_gain1.0000
6:116689319:G:GAacceptor_gain1.0000
6:116689319:GAT:Gacceptor_gain1.0000
6:116689319:GATG:Gacceptor_gain1.0000
6:116689449:AACAG:Adonor_gain1.0000
6:116689450:ACAG:Adonor_gain1.0000
6:116689451:CAG:Cdonor_gain1.0000
6:116689452:AG:Adonor_gain1.0000
6:116689452:AGGTA:Adonor_loss1.0000
6:116689453:GG:Gdonor_gain1.0000
6:116689454:G:GGdonor_gain1.0000
6:116689454:GTA:Gdonor_loss1.0000
6:116692053:A:AGacceptor_gain1.0000
6:116692054:G:GGacceptor_gain1.0000
6:116692054:GTT:Gacceptor_gain1.0000
6:116692154:GAG:Gdonor_gain1.0000
6:116692286:GTTTT:Gacceptor_loss1.0000
6:116692287:TTTTA:Tacceptor_loss1.0000
6:116692288:TTTA:Tacceptor_loss1.0000
6:116692289:TTAGG:Tacceptor_loss1.0000

AlphaMissense

3567 scored. Top likely-pathogenic:

dbSNP variants (sampled 300 via entrez): RS1000015104 (6:116730286 C>T), RS1000032614 (6:116718759 A>G), RS1000075617 (6:116716392 T>C), RS1000184899 (6:116715339 T>C), RS1000217661 (6:116703118 A>G), RS1000258892 (6:116679861 A>G), RS1000323134 (6:116709567 A>G), RS1000336771 (6:116737071 T>C), RS1000360917 (6:116709551 C>A), RS1000446896 (6:116703332 G>A), RS1000480008 (6:116703424 G>A,T), RS1000652410 (6:116709807 C>T), RS1000677750 (6:116741256 G>A), RS1000679012 (6:116698068 G>A), RS1000748021 (6:116696014 T>G)

Disease associations

OMIM: gene MIM:604545 | 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
GCST001024_3Telomere length3.000000e-06
GCST008991_7Early cardiac repolarization6.000000e-07

EFO canonical traits (1, from GWAS)

EFO IDTrait name
EFO:0004885early cardiac repolarization measurement

Drugs & pharmacology

Drug and pharmacology data

Is drug target: no

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

CTD chemical–gene interactions

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

ChemicalActions (top 5)PubMed papers
Valproic Acidaffects cotreatment, decreases expression6
bisphenol Aincreases expression, affects expression, affects methylation, affects cotreatment3
Benzo(a)pyrenedecreases expression, increases methylation3
trichostatin Aaffects cotreatment, decreases expression2
Tetrachlorodibenzodioxindecreases expression2
GSK-J4increases expression1
dicrotophosdecreases expression1
methylmercuric chloridedecreases expression1
triphenyl phosphateaffects expression1
tris(1,3-dichloro-2-propyl)phosphateincreases expression1
sodium arseniteincreases expression1
butyraldehydedecreases expression1
benzo(e)pyrenedecreases methylation1
aflatoxin B2decreases methylation1
di-n-butylphosphoric acidaffects expression1
CGP 52608affects binding, increases reaction1
4-(5-benzo(1,3)dioxol-5-yl-4-pyridin-2-yl-1H-imidazol-2-yl)benzamideaffects cotreatment, decreases expression1
nutlin 3affects cotreatment, increases secretion1
ICG 001decreases expression1
abrineincreases expression1
dorsomorphinaffects cotreatment, decreases expression1
jinfukangdecreases expression1
Resveratrolaffects cotreatment, increases expression1
Temozolomidedecreases expression1
Leflunomideincreases expression1
Cadmiumincreases abundance, increases expression1
Cisplatindecreases expression1
Dactinomycinaffects cotreatment, increases secretion1
Dexamethasoneaffects cotreatment, increases expression1
Doxorubicinincreases 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.