SEC14L2

gene
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Also known as TAPSPFKIAA1186KIAA1658TAP1

Summary

SEC14L2 (SEC14 like lipid binding 2, HGNC:10699) is a protein-coding gene on chromosome 22q12.2, encoding SEC14-like protein 2 (O76054). Carrier protein.

This gene encodes a cytosolic protein which belongs to a family of lipid-binding proteins including Sec14p, alpha-tocopherol transfer protein, and cellular retinol-binding protein. The encoded protein stimulates squalene monooxygenase which is a downstream enzyme in the cholesterol biosynthetic pathway. Alternatively spliced transcript variants encoding different isoforms have been identified for this gene.

Source: NCBI Gene 23541 — RefSeq curated summary.

At a glance

  • GWAS associations: 2
  • Clinical variants (ClinVar): 50 total
  • MANE Select transcript: NM_012429

Identifiers

Gene identifiers

FieldValue
HGNC IDHGNC:10699
Approved symbolSEC14L2
NameSEC14 like lipid binding 2
Location22q12.2
Locus typegene with protein product
StatusApproved
AliasesTAP, SPF, KIAA1186, KIAA1658, TAP1
Ensembl geneENSG00000100003
Ensembl biotypeprotein_coding
OMIM607558
Entrez23541

Gene structure

Transcript identifiers

Ensembl transcripts: 17 — 7 protein_coding, 4 protein_coding_CDS_not_defined, 4 retained_intron, 2 nonsense_mediated_decay

ENST00000402592, ENST00000405717, ENST00000415072, ENST00000416523, ENST00000428195, ENST00000429917, ENST00000437022, ENST00000452649, ENST00000459728, ENST00000464335, ENST00000483116, ENST00000484486, ENST00000485482, ENST00000615189, ENST00000617837, ENST00000619483, ENST00000620251

RefSeq mRNA: 4 — MANE Select: NM_012429 NM_001204204, NM_001291932, NM_012429, NM_033382

CCDS: CCDS13876, CCDS46685, CCDS56228

Canonical transcript exons

ENST00000615189 — 12 exons

ExonStartEnd
ENSE000034623073041594830416087
ENSE000036127473040942630409486
ENSE000036383883041623430416403
ENSE000036556603041059630410679
ENSE000036889733040918730409282
ENSE000037026443040741530407603
ENSE000037072623039964330399718
ENSE000037106233039701830397170
ENSE000037109443040709530407154
ENSE000037113193040634230406385
ENSE000037408093042227730425303
ENSE000037884543041575930415865

Expression profiles

Bgee: expression breadth ubiquitous, 207 present calls, max score 96.90.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 11.9195 / max 265.8515, expressed in 1524 samples.

FANTOM5 promoters (4 alternative TSS)

Promoter IDTPM avgSamples expressed
19167911.15371497
1916780.4635252
1916810.181991
1916800.120449

Top tissues by expression

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

TissueAnatomy IDExpression scoreQuality
right lobe of liverUBERON:000111496.90gold quality
lower esophagus mucosaUBERON:003583493.22gold quality
liverUBERON:000210791.99gold quality
right frontal lobeUBERON:000281090.76gold quality
prefrontal cortexUBERON:000045190.53gold quality
nucleus accumbensUBERON:000188290.41gold quality
amygdalaUBERON:000187689.88gold quality
C1 segment of cervical spinal cordUBERON:000646989.83gold quality
caudate nucleusUBERON:000187389.53gold quality
cingulate cortexUBERON:000302789.53gold quality
putamenUBERON:000187489.34gold quality
anterior cingulate cortexUBERON:000983589.32gold quality
pigmented layer of retinaUBERON:000178289.20gold quality
Brodmann (1909) area 9UBERON:001354088.10gold quality
esophagus mucosaUBERON:000246987.75gold quality
spinal cordUBERON:000224087.18gold quality
dorsolateral prefrontal cortexUBERON:000983487.11gold quality
hypothalamusUBERON:000189886.52gold quality
frontal cortexUBERON:000187086.49gold quality
neocortexUBERON:000195086.45gold quality
telencephalonUBERON:000189385.85gold quality
forebrainUBERON:000189085.28gold quality
cerebral cortexUBERON:000095684.88gold quality
corpus epididymisUBERON:000435984.76gold quality
substantia nigraUBERON:000203884.68gold quality
central nervous systemUBERON:000101784.28gold quality
brainUBERON:000095584.19gold quality
adenohypophysisUBERON:000219684.17gold quality
Ammon’s hornUBERON:000195484.02gold quality
sural nerveUBERON:001548883.92gold quality

Single-cell (SCXA)

Detected in 3 experiment(s), a significant marker in 1.

ExperimentMarker?Max mean expression
E-ANND-3yes4.13
E-ENAD-17no744.15
E-CURD-10no434.73

Regulation

Is transcription factor: yes

Downstream targets (CollecTRI)

1 targets.

TargetRegulation
PSMB9Unknown

miRNA regulators (miRDB)

62 targeting SEC14L2, 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-366299.9973.825684
HSA-MIR-569699.9872.364487
HSA-MIR-6778-3P99.9667.292693
HSA-MIR-345-3P99.8970.231421
HSA-MIR-629-3P99.8567.991875
HSA-MIR-431999.7669.832586
HSA-MIR-132-3P99.7370.561424
HSA-MIR-212-3P99.7370.651424
HSA-MIR-6752-3P99.7266.711587
HSA-MIR-120099.7170.421838
HSA-MIR-670-5P99.6769.941565
HSA-MIR-397599.6265.97697
HSA-MIR-1260A99.6166.671098
HSA-MIR-1260B99.6166.671098
HSA-MIR-1212399.5271.792990
HSA-MIR-183-3P99.4169.411598
HSA-MIR-125A-5P99.3670.591640
HSA-MIR-125B-5P99.3670.361662
HSA-MIR-6853-3P99.3670.791558
HSA-MIR-664A-3P99.2271.082696
HSA-MIR-544B99.1867.411632
HSA-MIR-429299.1665.571767
HSA-MIR-6791-5P99.1665.921844
HSA-MIR-6734-3P99.1566.271627
HSA-MIR-432499.0470.141569
HSA-MIR-4650-3P99.0168.391062
HSA-MIR-314298.8866.09529
HSA-MIR-1245B-5P98.8866.55576
HSA-MIR-6760-5P98.8766.731515
HSA-MIR-4477A98.8369.752952

Literature-anchored findings (GeneRIF, showing 18)

  • SPF has two domain topology, and the ligand binding cavity has a peculiar horseshoe-like shape (PMID:12429094)
  • phosphorylation enhances its ability to stimulate microsomal squalene monooxygenase; phosphorylation may provide a means for the rapid short term modulation of cholesterol synthesis (PMID:12454003)
  • Data report the three-dimensional crystal structure of human supernatant protein factor in complex with RRR-alpha-tocopherylquinone, the major physiological oxidation product of RRR-alpha-tocopherol, at a resolution of 1.95A. (PMID:12972248)
  • Discusses cloning of rat SEC14-like protein 3 and mentions three human SPF-related genes, known as SEC14-like protein 1, 2 and 3. (PMID:15033454)
  • TAP not only mediates vitamin E absorption to facilitate vitamin E antiproliferation effect in prostate cancer cells, but also functions like a tumor suppressor gene to control cancer cell viability through a non-vitamin E manner. (PMID:16267002)
  • reduced expression of TAP was associated with the cell proliferation status of prostate cancer, adverse pathological parameters and the increased risk of recurrence (PMID:17334589)
  • Genetic variation in TTPA and SEC14L2 is associated with serum alpha-tocopherol but does not have a direct effect on prostate cancer when vitamin E is administered. (PMID:19190344)
  • These findings are consistent with alpha-tocopherol-associated protein acting as an antiproliferative factor in estrogen-receptor-positive luminal cells in normal/benign breast tissue (PMID:19305383)
  • Fndings raise the possibility that TAP/Sec14L2 may serve as a tumor suppressor in breast carcinogenesis. (PMID:19909011)
  • novel association between the rs737723 polymorphism (SEC14L2/TAP) and higher primary open-angle glaucoma risk (PMID:23401652)
  • TAP expression was significantly correlated with Her2/neu receptor expression and breast cancer stage. (PMID:23411208)
  • hTAP1 reduces the in vitro activity of the phosphatidylinositol-3-kinase gamma (PI3Kgamma) indicating the formation of a stalled/inactive hTAP1/PI3Kgamma heterodimer. (PMID:24983950)
  • PI3Kgamma-induced VEGF expression was reduced when the human tocopherol-associated protein 1 (hTAP1/SEC14L2) was overexpressed suggesting formation of an inactive PI3Kgamma/hTAP1 heterodimer. (PMID:25290554)
  • SEC14L2 enabled RNA replication of diverse HCV genotypes in several human hepatoma cell lines (PMID:26266980)
  • Here, the authors report that the zebrafish/human phosphatidylinositol transfer protein Sec14l3/SEC14L2 act as GTPase proteins to transduce Wnt signals from Frizzled to phospholipase C (PLC). (PMID:28463110)
  • SEC14L2, a lipid-binding protein, regulates HCV replication in culture with inter- and intra-genotype variations. (PMID:30472319)
  • Discovering master regulators in hepatocellular carcinoma: one novel MR, SEC14L2 inhibits cancer cells. (PMID:31851620)
  • TAP-ing into the cross-presentation secrets of dendritic cells. (PMID:37116384)

Cross-species orthologs

9 orthologs

OrganismSymbolGene ID
danio_reriosec14l7ENSDARG00000099633
mus_musculusSec14l2ENSMUSG00000003585
rattus_norvegicusSec14l2ENSRNOG00000004672
drosophila_melanogasterretmFBGN0031814
drosophila_melanogasterCG13893FBGN0035146
caenorhabditis_elegansWBGENE00007925
caenorhabditis_elegansWBGENE00009241
caenorhabditis_elegansctg-1WBGENE00010370
caenorhabditis_elegansWBGENE00011962

Paralogs (6): SEC14L3 (ENSG00000100012), SEC14L5 (ENSG00000103184), SEC14L1 (ENSG00000129657), SEC14L4 (ENSG00000133488), TTPA (ENSG00000137561), SEC14L6 (ENSG00000214491)

Protein

Protein identifiers

SEC14-like protein 2O76054 (reviewed: O76054)

Alternative names: Alpha-tocopherol-associated protein, Squalene transfer protein, Supernatant protein factor

All UniProt accessions (7): O76054, B3KRD8, C9JTM4, C9JYY7, C9JZI9, F8WED8, F8WEE7

UniProt curated annotations — full annotation on UniProt →

Function. Carrier protein. Binds to some hydrophobic molecules and promotes their transfer between the different cellular sites. Binds with high affinity to alpha-tocopherol. Also binds with a weaker affinity to other tocopherols and to tocotrienols. May have a transcriptional activatory activity via its association with alpha-tocopherol. Probably recognizes and binds some squalene structure, suggesting that it may regulate cholesterol biosynthesis by increasing the transfer of squalene to a metabolic active pool in the cell.

Subunit / interactions. Monomer.

Subcellular location. Cytoplasm. Nucleus.

Tissue specificity. Widely expressed. Strong expression in liver, brain and prostate.

Isoforms (3)

UniProt IDNamesCanonical?
O76054-11yes
O76054-42
O76054-53

RefSeq proteins (4): NP_001191133, NP_001278861, NP_036561, NP_203740 (=MANE)

Domains & families (InterPro)

IDNameType
IPR001251CRAL-TRIO_domDomain
IPR009038GOLD_domDomain
IPR011074CRAL/TRIO_N_domDomain
IPR036273CRAL/TRIO_N_dom_sfHomologous_superfamily
IPR036598GOLD_dom_sfHomologous_superfamily
IPR036865CRAL-TRIO_dom_sfHomologous_superfamily
IPR051064

Pfam: PF00650, PF03765

UniProt features (50 total): helix 19, strand 15, turn 5, modified residue 4, domain 2, splice variant 2, chain 1, sequence conflict 1, sequence variant 1

Structure

Experimental structures (PDB)

4 structures.

PDBMethodResolution (Å)
4OMJX-RAY DIFFRACTION1.6
4OMKX-RAY DIFFRACTION1.75
1OLMX-RAY DIFFRACTION1.95
1O6UX-RAY DIFFRACTION2.05

Predicted structure (AlphaFold)

ModelpLDDTFraction very-high
AF-O76054-F195.760.93

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 (4): 51, 253, 257, 393

Function

Pathways and Gene Ontology

Reactome pathways

0 pathways

MSigDB gene sets: 630 (showing top): WALLACE_PROSTATE_CANCER_RACE_UP, BENPORATH_ES_WITH_H3K27ME3, REACTOME_ADAPTIVE_IMMUNE_SYSTEM, REACTOME_ANTIGEN_PRESENTATION_FOLDING_ASSEMBLY_AND_PEPTIDE_LOADING_OF_CLASS_I_MHC, REACTOME_CLASS_I_MHC_MEDIATED_ANTIGEN_PROCESSING_PRESENTATION, MODULE_45, GRAESSMANN_APOPTOSIS_BY_DOXORUBICIN_UP, GOBP_ANTIGEN_PROCESSING_AND_PRESENTATION_OF_PEPTIDE_ANTIGEN, BROWNE_HCMV_INFECTION_12HR_UP, KEGG_ABC_TRANSPORTERS, NFKB_Q6, HINATA_NFKB_TARGETS_KERATINOCYTE_UP, GOBP_REGULATION_OF_LIPID_BIOSYNTHETIC_PROCESS, GOBP_REGULATION_OF_CHOLESTEROL_BIOSYNTHETIC_PROCESS, BENNETT_SYSTEMIC_LUPUS_ERYTHEMATOSUS

GO Biological Process (2): regulation of cholesterol biosynthetic process (GO:0045540), positive regulation of DNA-templated transcription (GO:0045893)

GO Molecular Function (3): phospholipid binding (GO:0005543), vitamin E binding (GO:0008431), lipid binding (GO:0008289)

GO Cellular Component (5): nucleus (GO:0005634), nucleoplasm (GO:0005654), cytoplasm (GO:0005737), cytosol (GO:0005829), extracellular exosome (GO:0070062)

GO top-level categories

Rollup of top GO terms by namespace:

CategoryTerms
cellular anatomical structure3
cholesterol biosynthetic process1
regulation of cholesterol metabolic process1
regulation of sterol biosynthetic process1
regulation of alcohol biosynthetic process1
DNA-templated transcription1
regulation of DNA-templated transcription1
positive regulation of RNA biosynthetic process1
lipid binding1
vitamin binding1
heterocyclic compound binding1
binding1
intracellular membrane-bounded organelle1
nuclear lumen1
intracellular anatomical structure1
cytoplasm1
extracellular vesicle1

Protein interactions and networks

STRING

572 interactions, top by confidence (×1000):

Protein AProtein BPartner UniProtScore
SEC14L2PI4KAP42356528
SEC14L2CD81P18582507
SEC14L2CLDN1O95832497
SEC14L2PPIAP05092470
SEC14L2SCARB1Q8WTV0461
SEC14L2DVL2O14641432
SEC14L2APOEP02649415
SEC14L2CD302Q8IX05379
SEC14L2SQLEQ14534361
SEC14L2CCDC74AQ96AQ1355
SEC14L2SRFBP1Q8NEF9349
SEC14L2CYP4F2P78329312
SEC14L2CCDC74BQ96LY2308
SEC14L2BUD13Q9BRD0307
SEC14L2LPLP06858292

IntAct

5 interactions, top by confidence:

ABTypeScore
SEC14L3SEC14L2psi-mi:“MI:0914”(association)0.530
SEC14L2Ppm1lpsi-mi:“MI:0915”(physical association)0.400
SPANXN2ZNF320psi-mi:“MI:0914”(association)0.350

BioGRID (28): SEC14L2 (Affinity Capture-MS), SEC14L2 (Affinity Capture-RNA), SEC14L2 (Affinity Capture-RNA), SEC14L2 (Affinity Capture-MS), SEC14L2 (Affinity Capture-MS), SEC14L2 (Co-fractionation), SEC14L2 (Co-fractionation), GOT2 (Co-fractionation), CSTB (Co-fractionation), GPI (Co-fractionation), RAC2 (Co-fractionation), ANXA7 (Co-fractionation), ALDH4A1 (Co-fractionation), PEBP1 (Co-fractionation), S100A11 (Co-fractionation)

ESM2 similar proteins: A7T167, B3MZN7, B4PYH5, B5MCN3, F4JYJ3, O17907, O35239, O76054, P24280, P24859, P34913, P34914, P43378, P49193, P53989, P56523, P58875, P80299, Q03606, Q16KN5, Q16P87, Q29JQ0, Q2TAF8, Q4V8Q1, Q5RFR0, Q641Z2, Q653S9, Q6IPT4, Q6J756, Q6Q2C2, Q75DK1, Q78EJ9, Q8L7L1, Q8R0F9, Q91VA3, Q99J08, Q99MS0, Q9BTX7, Q9D3D0, Q9FKS0

Diamond homologs: A8Y5H7, B5MCN3, F4IHJ0, F4J7S8, F4JLE5, F4JVA6, F4JVA9, F4JYJ3, F4K6D3, O43304, O76054, P49193, P58875, Q03606, Q0V9N0, Q16KN5, Q29JQ0, Q7PWB1, Q8GXC6, Q8R0F9, Q92503, Q93ZE9, Q94A34, Q99J08, Q99MS0, Q9SI13, Q9SIW3, Q9UDX3, Q9UDX4, Q9VMD6, Q9Z1J8, Q10137, F4HP88, P24280, P24859, P33324, P45816, P46250, P53989, Q501H5

SIGNOR signaling

3 interactions.

AEffectBMechanism
PKA“up-regulates activity”SEC14L2phosphorylation
PRKACA“up-regulates activity”SEC14L2phosphorylation

Disease & clinical

Clinical variants and AI predictions

ClinVar

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

ClassificationCount (floor)
Pathogenic0
Likely pathogenic0
Uncertain significance36
Likely benign2
Benign1

Top pathogenic / likely-pathogenic (0)

SpliceAI

3289 predictions. Top by Δscore:

VariantEffectΔscore
22:30397168:AAGG:Adonor_loss1.0000
22:30397169:AGGTG:Adonor_loss1.0000
22:30397170:GGT:Gdonor_loss1.0000
22:30397171:GTGAG:Gdonor_loss1.0000
22:30397172:T:Adonor_loss1.0000
22:30399641:A:AGacceptor_gain1.0000
22:30399642:G:GGacceptor_gain1.0000
22:30399725:G:Tdonor_gain1.0000
22:30406340:A:AGacceptor_gain1.0000
22:30406341:G:GGacceptor_gain1.0000
22:30406384:AGGT:Adonor_loss1.0000
22:30406386:GTGAG:Gdonor_loss1.0000
22:30406387:T:Gdonor_loss1.0000
22:30407093:A:AGacceptor_gain1.0000
22:30407094:G:GGacceptor_gain1.0000
22:30407094:GCAT:Gacceptor_gain1.0000
22:30407413:A:AGacceptor_gain1.0000
22:30407413:AG:Aacceptor_gain1.0000
22:30407413:AGGT:Aacceptor_gain1.0000
22:30407414:G:Aacceptor_loss1.0000
22:30407414:G:GGacceptor_gain1.0000
22:30407414:GG:Gacceptor_gain1.0000
22:30407414:GGT:Gacceptor_gain1.0000
22:30407414:GGTG:Gacceptor_gain1.0000
22:30407414:GGTGA:Gacceptor_gain1.0000
22:30407599:CAAAG:Cdonor_gain1.0000
22:30407600:AAAG:Adonor_gain1.0000
22:30407601:AAG:Adonor_gain1.0000
22:30407601:AAGGT:Adonor_loss1.0000
22:30407602:AG:Adonor_gain1.0000

AlphaMissense

2655 scored. Top likely-pathogenic:

VariantProtein changeam_pathogenicity
22:30410617:C:AA201D0.999
22:30416235:T:AW305R0.999
22:30416235:T:CW305R0.999
22:30416242:T:CF307S0.999
22:30422289:T:CF365S0.999
22:30422296:C:AN367K0.999
22:30422296:C:GN367K0.999
22:30399709:T:AW41R0.998
22:30399709:T:CW41R0.998
22:30407473:T:AV98D0.998
22:30410626:T:CL204P0.998
22:30416084:T:AL303H0.998
22:30416084:T:CL303P0.998
22:30416271:G:TG317W0.998
22:30416272:G:AG317E0.998
22:30416320:T:CM333T0.998
22:30416387:C:GC355W0.998
22:30422280:T:AV362D0.998
22:30422286:G:CR364P0.998
22:30422303:A:CS370R0.998
22:30422305:C:AS370R0.998
22:30422305:C:GS370R0.998
22:30422320:G:CK375N0.998
22:30422320:G:TK375N0.998
22:30422325:T:AV377D0.998
22:30407508:G:CG110R0.997
22:30409253:T:AW164R0.997
22:30409253:T:CW164R0.997
22:30409478:T:AV191D0.997
22:30410614:T:AV200E0.997

dbSNP variants (sampled 300 via entrez): RS1000135396 (22:30414019 C>T), RS1000235419 (22:30397460 C>CA), RS1000241369 (22:30421813 C>G), RS1000696529 (22:30424827 G>T), RS1000726765 (22:30397071 C>A,T), RS1000744980 (22:30401523 AT>A,ATT), RS1000783329 (22:30408428 CA>C,CAA), RS1000943325 (22:30410043 T>C), RS1000964699 (22:30408050 C>T), RS1001175892 (22:30402851 G>A), RS1001206675 (22:30413939 A>G), RS1001231908 (22:30396868 C>T), RS1001394522 (22:30395430 A>G), RS1001445670 (22:30399845 C>G), RS1001498220 (22:30396792 A>G)

Disease associations

OMIM: gene MIM:607558 | 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
GCST001585_24Breast size4.000000e-07
GCST004607_193Plateletcrit7.000000e-09

EFO canonical traits (1, from GWAS)

EFO IDTrait name
EFO:0007985platelet crit

Drugs & pharmacology

Drug and pharmacology data

Is drug target: no

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

PharmGKB variants

2 variants.

VariantGenesLevelScore#Clin annotsDrugs
rs2071543PSMB8, TAP1, TAP20.000
rs1135216TAP10.000

CTD chemical–gene interactions

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

ChemicalActions (top 5)PubMed papers
Benzo(a)pyrenedecreases expression, increases methylation3
Valproic Acidaffects cotreatment, decreases expression, increases expression3
Estradiolaffects cotreatment, increases expression2
Smokeincreases abundance, decreases expression2
Testosteroneincreases expression, affects cotreatment, decreases reaction2
Tobacco Smoke Pollutionaffects expression, increases expression2
aristolochic acid Iincreases expression1
sotorasibaffects cotreatment, decreases expression1
dicrotophosdecreases expression1
pirinixic acidincreases activity, affects binding, decreases expression1
bisphenol Adecreases expression1
terbufosaffects response to substance1
alpha-terpineoldecreases reaction, increases expression1
sodium arsenitedecreases expression1
linalooldecreases reaction, increases expression1
gamma-terpineneincreases expression, decreases reaction1
potassium chromate(VI)decreases expression1
terpinenol-4decreases reaction, increases expression1
lavender oildecreases reaction, increases expression1
di-n-butylphosphoric acidaffects expression1
linalyl acetateincreases expression, decreases reaction1
perfluoro-n-nonanoic aciddecreases expression1
4-(5-benzo(1,3)dioxol-5-yl-4-pyridin-2-yl-1H-imidazol-2-yl)benzamideaffects cotreatment, decreases expression1
dorsomorphinaffects cotreatment, decreases expression1
trametinibaffects cotreatment, decreases expression1
NVP-BKM120affects cotreatment, decreases expression1
Limonenedecreases reaction, increases expression1
Eucalyptoldecreases reaction, increases expression1
Air Pollutantsdecreases expression, increases abundance1
Calcitriolincreases expression, affects cotreatment1

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.