PUM3

gene
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Also known as XTP5PENPUF6hPUF-AHA-8Puf-A

Summary

PUM3 (pumilio RNA binding family member 3, HGNC:29676) is a protein-coding gene on chromosome 9p24.2, encoding Pumilio homolog 3 (Q15397). Inhibits the poly(ADP-ribosyl)ation activity of PARP1 and the degradation of PARP1 by CASP3 following genotoxic stress.

Enables RNA binding activity. Predicted to be involved in regulation of translation. Located in chromosome; endoplasmic reticulum; and nuclear lumen.

Source: NCBI Gene 9933 — RefSeq curated summary.

At a glance

  • GWAS associations: 9
  • Clinical variants (ClinVar): 151 total — 1 pathogenic
  • MANE Select transcript: NM_014878

Identifiers

Gene identifiers

FieldValue
HGNC IDHGNC:29676
Approved symbolPUM3
Namepumilio RNA binding family member 3
Location9p24.2
Locus typegene with protein product
StatusApproved
AliasesXTP5, PEN, PUF6, hPUF-A, HA-8, Puf-A
Ensembl geneENSG00000080608
Ensembl biotypeprotein_coding
OMIM609960
Entrez9933

Gene structure

Transcript identifiers

Ensembl transcripts: 29 — 26 protein_coding, 1 retained_intron, 1 protein_coding_CDS_not_defined, 1 nonsense_mediated_decay

ENST00000382032, ENST00000397885, ENST00000469168, ENST00000490444, ENST00000861028, ENST00000861029, ENST00000861030, ENST00000861031, ENST00000861032, ENST00000861033, ENST00000861034, ENST00000861035, ENST00000861036, ENST00000922205, ENST00000922206, ENST00000922207, ENST00000922208, ENST00000922209, ENST00000922210, ENST00000922211, ENST00000922212, ENST00000922213, ENST00000922214, ENST00000922215, ENST00000922216, ENST00000922217, ENST00000954334, ENST00000954335, ENST00000954336

RefSeq mRNA: 1 — MANE Select: NM_014878 NM_014878

CCDS: CCDS6448

Canonical transcript exons

ENST00000397885 — 18 exons

ExonStartEnd
ENSE0000068673028103442810431
ENSE0000081308728078142807904
ENSE0000098207928312512831344
ENSE0000098208028309622831028
ENSE0000098208128297742829948
ENSE0000098208228286752828778
ENSE0000098208328270732827151
ENSE0000098208528237812823834
ENSE0000098208628200182820098
ENSE0000098208728122202812362
ENSE0000098208828113612811583
ENSE0000102834528333572833432
ENSE0000108895228247172824815
ENSE0000153066528440452844095
ENSE0000159651828371802837401
ENSE0000160382728340312834166
ENSE0000178480628384262838517
ENSE0000191428628041522804463

Expression profiles

Bgee: expression breadth ubiquitous, 264 present calls, max score 94.70.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 26.1351 / max 875.5705, expressed in 1777 samples.

FANTOM5 promoters (1 alternative TSS)

Promoter IDTPM avgSamples expressed
9974926.13511777

Top tissues by expression

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

TissueAnatomy IDExpression scoreQuality
adrenal tissueUBERON:001830394.70gold quality
calcaneal tendonUBERON:000370192.83gold quality
mucosa of transverse colonUBERON:000499192.29gold quality
esophagus mucosaUBERON:000246992.26gold quality
lower esophagus mucosaUBERON:003583491.85gold quality
primordial germ cell in gonadCL:0000670 ∩ UBERON:000099191.83gold quality
metanephros cortexUBERON:001053391.45gold quality
peritoneumUBERON:000235891.31gold quality
omental fat padUBERON:001041491.31gold quality
male germ line stem cell (sensu Vertebrata) in testisCL:0000089 ∩ UBERON:000047391.04gold quality
olfactory segment of nasal mucosaUBERON:000538691.04gold quality
left ovaryUBERON:000211991.00gold quality
adipose tissue of abdominal regionUBERON:000780890.62gold quality
minor salivary glandUBERON:000183090.60gold quality
body of pancreasUBERON:000115090.44gold quality
right ovaryUBERON:000211890.38gold quality
upper lobe of left lungUBERON:000895290.22gold quality
left uterine tubeUBERON:000130390.20gold quality
mouth mucosaUBERON:000372989.96gold quality
rectumUBERON:000105289.85gold quality
esophagusUBERON:000104389.75gold quality
ventricular zoneUBERON:000305389.67gold quality
skin of abdomenUBERON:000141689.54gold quality
tonsilUBERON:000237289.49gold quality
sural nerveUBERON:001548889.43gold quality
left adrenal gland cortexUBERON:003582589.39gold quality
upper lobe of lungUBERON:000894889.38gold quality
ovaryUBERON:000099289.28gold quality
islet of LangerhansUBERON:000000689.11gold quality
right adrenal glandUBERON:000123389.05gold quality

Single-cell (SCXA)

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

ExperimentMarker?Max mean expression
E-MTAB-4850no3179.70
E-MTAB-6142no218.07
E-ANND-3no0.00

Regulation

Is transcription factor: no

miRNA regulators (miRDB)

21 targeting PUM3, 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-450A-1-3P100.0069.331837
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-520D-5P99.9873.344883
HSA-MIR-524-5P99.9873.434882
HSA-MIR-627-3P99.9071.423316
HSA-MIR-391999.8769.452489
HSA-MIR-544A99.8468.661965
HSA-MIR-3121-3P99.8271.963630
HSA-MIR-6739-5P99.8067.872806
HSA-MIR-4668-5P99.7970.583782
HSA-MIR-6733-5P99.7467.942759
HSA-MIR-315399.5567.592337
HSA-MIR-608399.4768.732393
HSA-MIR-410-3P99.2769.982457
HSA-MIR-3925-5P99.2167.901466
HSA-MIR-477398.3567.301710
HSA-MIR-6502-3P97.8665.43569

Literature-anchored findings (GeneRIF, showing 2)

  • Results indicate that mismatches in minor H antigens HA-8 (KIAA0020) and ACC-1 (BCL2A1) predisposed to chronic graft-versus-host disease (GvHD). (PMID:23480177)
  • Phosphorylation of PUF-A/PUM3 on Y259 modulates PUF-A stability and cell proliferation. (PMID:34407138)

Cross-species orthologs

6 orthologs

OrganismSymbolGene ID
danio_reriopum3ENSDARG00000087779
mus_musculusPum3ENSMUSG00000041360
rattus_norvegicusPum3ENSRNOG00000012574
rattus_norvegicusPum3l1ENSRNOG00000048356
drosophila_melanogasterpengFBGN0015527
caenorhabditis_elegansWBGENE00014165

Protein

Protein identifiers

Pumilio homolog 3Q15397 (reviewed: Q15397)

Alternative names: HBV X-transactivated gene 5 protein, HBV XAg-transactivated protein 5, Minor histocompatibility antigen HA-8

All UniProt accessions (2): Q15397, S4R3K8

UniProt curated annotations — full annotation on UniProt →

Function. Inhibits the poly(ADP-ribosyl)ation activity of PARP1 and the degradation of PARP1 by CASP3 following genotoxic stress. Binds to double-stranded RNA or DNA without sequence specificity. Involved in development of the eye and of primordial germ cells.

Subunit / interactions. Interacts with PARP1 (via catalytic domain).

Subcellular location. Nucleus. Nucleolus. Nucleoplasm. Chromosome.

Tissue specificity. Widely expressed.

Domain organisation. The HA-8 region can be cleaved and exposed at the cell surface where it plays a role as a minor histocompatibility HLA-A*0201-restricted antigen. A 90 degree bend between Pumilio repeats 3 and 4 gives rise to a L-shaped protein.

Polymorphism. The following alleles of HA-8 are known: HA-8R, HA-8P, HA-8PL, of which only HA-8R leads to specific cytotoxic T lymphocyte (CTL) recognition. The lack of CTL recognition of cells expressing HA-8P may be due to impaired transport associated with antigen processing. The sequence shown is that of HA-8R.

RefSeq proteins (1): NP_055693* (*=MANE)

Domains & families (InterPro)

IDNameType
IPR001313Pumilio_RNA-bd_rptRepeat
IPR011989ARM-likeHomologous_superfamily
IPR012959CPL_domDomain
IPR016024ARM-type_foldHomologous_superfamily
IPR033133PUM-HDDomain
IPR040059PUM3Family

Pfam: PF08144

UniProt features (82 total): helix 45, repeat 11, turn 8, sequence variant 6, compositionally biased region 3, region of interest 2, strand 2, chain 1, domain 1, short sequence motif 1, modified residue 1, sequence conflict 1

Structure

Experimental structures (PDB)

2 structures.

PDBMethodResolution (Å)
4WZRX-RAY DIFFRACTION2.15
4WZWX-RAY DIFFRACTION2.95

Predicted structure (AlphaFold)

ModelpLDDTFraction very-high
AF-Q15397-F185.190.76

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 (1): 33

Function

Pathways and Gene Ontology

Reactome pathways

0 pathways

MSigDB gene sets: 203 (showing top): RODRIGUES_THYROID_CARCINOMA_ANAPLASTIC_UP, GRAESSMANN_APOPTOSIS_BY_DOXORUBICIN_DN, GRAESSMANN_RESPONSE_TO_MC_AND_DOXORUBICIN_DN, GOBP_REGULATION_OF_TRANSFERASE_ACTIVITY, TAKADA_GASTRIC_CANCER_COPY_NUMBER_DN, PUJANA_CHEK2_PCC_NETWORK, GOBP_TRANSLATION, WEI_MYCN_TARGETS_WITH_E_BOX, GOBP_POST_TRANSCRIPTIONAL_REGULATION_OF_GENE_EXPRESSION, SCHUHMACHER_MYC_TARGETS_UP, ONKEN_UVEAL_MELANOMA_UP, MYOD_01, MILI_PSEUDOPODIA_HAPTOTAXIS_UP, YANG_BREAST_CANCER_ESR1_DN, BILD_E2F3_ONCOGENIC_SIGNATURE

GO Biological Process (2): regulation of translation (GO:0006417), regulation of protein ADP-ribosylation (GO:0010835)

GO Molecular Function (4): DNA binding (GO:0003677), RNA binding (GO:0003723), mRNA binding (GO:0003729), protein binding (GO:0005515)

GO Cellular Component (5): nucleoplasm (GO:0005654), chromosome (GO:0005694), nucleolus (GO:0005730), endoplasmic reticulum (GO:0005783), nucleus (GO:0005634)

GO top-level categories

Rollup of top GO terms by namespace:

CategoryTerms
nucleic acid binding2
nuclear lumen2
intracellular membraneless organelle2
intracellular membrane-bounded organelle2
translation1
post-transcriptional regulation of gene expression1
regulation of protein metabolic process1
regulation of transferase activity1
NAD+-protein mono-ADP-ribosyltransferase activity1
RNA binding1
binding1
cellular anatomical structure1
cytoplasm1
endomembrane system1

Protein interactions and networks

STRING

2098 interactions, top by confidence (×1000):

Protein AProtein BPartner UniProtScore
PUM3EIF5BO60841723
PUM3NOP9Q86U38678
PUM3HLA-AP01891678
PUM3NME2P22392621
PUM3NSA2O95478562
PUM3RPF2Q9H7B2526
PUM3PUM1Q14671515
PUM3PUM2Q8TB72500
PUM3DDX27Q96GQ7475
PUM3ARHGAP45Q92619470
PUM3COX17Q14061462
PUM3SURF6O75683455
PUM3GNL2Q13823452
PUM3TMEM209Q96SK2443
PUM3PNO1Q9NRX1440

IntAct

256 interactions, top by confidence:

ABTypeScore
MED23MED19psi-mi:“MI:2364”(proximity)0.770
SRP68SRP72psi-mi:“MI:0914”(association)0.730
RPL10ARRP8psi-mi:“MI:0914”(association)0.640
RPL14RRP8psi-mi:“MI:0914”(association)0.640
H1-1RRP8psi-mi:“MI:0914”(association)0.640
NOP53RRP8psi-mi:“MI:0914”(association)0.640
NOL12RRP8psi-mi:“MI:0914”(association)0.640
NPM1NVLpsi-mi:“MI:0914”(association)0.610
MAGEB10GTPBP10psi-mi:“MI:0914”(association)0.530
H1-6ZNF724psi-mi:“MI:0914”(association)0.530
RBM34NVLpsi-mi:“MI:0914”(association)0.530
H1-4IGF2BP3psi-mi:“MI:0914”(association)0.530
ILF2IGF2BP3psi-mi:“MI:0914”(association)0.530
RPL37AMPHOSPH10psi-mi:“MI:0914”(association)0.530
ZNF2MPHOSPH10psi-mi:“MI:0914”(association)0.530
RPL18NOP56psi-mi:“MI:0914”(association)0.530
MACROH2A2PPM1Gpsi-mi:“MI:0914”(association)0.530
PRR11NVLpsi-mi:“MI:0914”(association)0.530
MAK16NVLpsi-mi:“MI:0914”(association)0.530
ZNF71NVLpsi-mi:“MI:0914”(association)0.530
RPL30RRP8psi-mi:“MI:0914”(association)0.530
RPL8RRP8psi-mi:“MI:0914”(association)0.530
RPL18ARRP8psi-mi:“MI:0914”(association)0.530
PUM3RRP8psi-mi:“MI:0914”(association)0.530
PDGFBDKC1psi-mi:“MI:0914”(association)0.530
NRBM47psi-mi:“MI:0914”(association)0.530

BioGRID (392): KIAA0020 (Affinity Capture-RNA), KIAA0020 (Affinity Capture-RNA), KIAA0020 (Affinity Capture-RNA), KIAA0020 (Affinity Capture-MS), KIAA0020 (Affinity Capture-MS), KIAA0020 (Affinity Capture-MS), KIAA0020 (Affinity Capture-MS), KIAA0020 (Affinity Capture-MS), KIAA0020 (Affinity Capture-MS), KIAA0020 (Affinity Capture-MS), KIAA0020 (Affinity Capture-MS), KIAA0020 (Affinity Capture-MS), KIAA0020 (Affinity Capture-MS), KIAA0020 (Affinity Capture-MS), KIAA0020 (Affinity Capture-MS)

ESM2 similar proteins: A1Z3X3, A2AT37, A4GWN3, A5PKE4, B0UYI1, B2GV38, O08848, O35841, O43395, O75031, P49754, Q08CZ0, Q15397, Q15650, Q2HJ41, Q2T9K6, Q3UBZ5, Q5EAQ1, Q5KU39, Q5R5F1, Q5R644, Q5RCR8, Q5RE03, Q5U3V9, Q5VZK9, Q5XIC7, Q5ZJ85, Q5ZKG8, Q5ZMW3, Q676U5, Q6GLK9, Q6GMH0, Q6NYU2, Q7Z3V4, Q80UM3, Q80YQ8, Q8BHL5, Q8BM39, Q8R2M0, Q8WV92

Diamond homologs: Q09622, Q15397, Q562C7, Q9LRZ3, Q8BKS9, X1WGX5, Q04373, Q9UU76

SIGNOR signaling

1 interactions.

AEffectBMechanism
“Cap-binding complex”“down-regulates quantity by repression”PUM3“post transcriptional regulation”

Enriched among interaction partners

Reactome pathways and GO biological processes over-represented among this gene’s 188 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
Formation of Senescence-Associated Heterochromatin Foci (SAHF)635.4×3e-07
Peptide chain elongation2426.7×3e-26
Viral mRNA Translation2426.7×3e-26
PELO:HBS1L and ABCE1 dissociate a ribosome on a non-stop mRNA2426.4×3e-26
Selenocysteine synthesis2425.3×6e-26
Eukaryotic Translation Termination2425.3×6e-26
Nonsense Mediated Decay (NMD) independent of the Exon Junction Complex (EJC)2424.8×9e-26
ZNF598 and the Ribosome-associated Quality Trigger (RQT) complex dissociate a ribosome stalled on a no-go mRNA2424.8×9e-26

GO biological processes:

GO termPartnersFoldFDR
negative regulation of DNA recombination747.1×1e-08
chromosome condensation735.3×8e-08
maturation of LSU-rRNA from tricistronic rRNA transcript (SSU-rRNA, 5.8S rRNA, LSU-rRNA)528.0×5e-05
cytoplasmic translation2527.7×9e-27
ribosomal large subunit biogenesis821.2×3e-07
positive regulation of viral genome replication517.4×4e-04
translation2515.4×4e-20
rRNA processing1815.3×4e-14

Disease & clinical

Clinical variants and AI predictions

ClinVar

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

ClassificationCount (floor)
Pathogenic1
Likely pathogenic0
Uncertain significance117
Likely benign3
Benign3

Top pathogenic / likely-pathogenic (1)

Variant IDHGVSClassification
563675GRCh37/hg19 9p24.3-23(chr9:203861-9306658)x1Pathogenic

SpliceAI

2698 predictions. Top by Δscore:

VariantEffectΔscore
9:2804464:C:CCacceptor_gain1.0000
9:2806445:T:Cacceptor_gain1.0000
9:2811605:G:GCacceptor_gain1.0000
9:2811608:T:TCacceptor_gain1.0000
9:2811624:C:CTacceptor_gain1.0000
9:2823836:T:Cacceptor_gain1.0000
9:2827071:A:ACdonor_gain1.0000
9:2827072:C:CCdonor_gain1.0000
9:2827072:CTGAT:Cdonor_gain1.0000
9:2828673:A:ACdonor_gain1.0000
9:2828674:C:CCdonor_gain1.0000
9:2828779:C:CCacceptor_gain1.0000
9:2829760:T:TAdonor_gain1.0000
9:2829770:TCAC:Tdonor_loss1.0000
9:2829771:CACCT:Cdonor_loss1.0000
9:2829808:T:TAdonor_gain1.0000
9:2829944:TACTT:Tacceptor_gain1.0000
9:2829946:CTT:Cacceptor_gain1.0000
9:2829946:CTTC:Cacceptor_loss1.0000
9:2829947:TT:Tacceptor_gain1.0000
9:2829947:TTCTA:Tacceptor_loss1.0000
9:2829948:TC:Tacceptor_loss1.0000
9:2829949:C:Aacceptor_loss1.0000
9:2829949:C:CCacceptor_gain1.0000
9:2829950:T:Gacceptor_loss1.0000
9:2829954:C:CTacceptor_gain1.0000
9:2829956:C:CTacceptor_gain1.0000
9:2829957:A:Tacceptor_gain1.0000
9:2829959:C:CTacceptor_gain1.0000
9:2829961:C:CTacceptor_gain1.0000

AlphaMissense

4303 scored. Top likely-pathogenic:

VariantProtein changeam_pathogenicity
9:2823826:T:AK381N0.999
9:2823826:T:GK381N0.999
9:2823829:C:AR380S0.999
9:2823829:C:GR380S0.999
9:2824717:C:AK378N0.999
9:2824717:C:GK378N0.999
9:2823827:T:AK381I0.998
9:2823828:T:CK381E0.998
9:2823830:C:AR380M0.998
9:2823830:C:GR380T0.998
9:2831320:G:TR181S0.998
9:2834034:C:GR146T0.998
9:2823814:T:AK385N0.997
9:2823814:T:GK385N0.997
9:2823815:T:AK385I0.997
9:2823816:T:CK385E0.997
9:2824727:C:TG375D0.997
9:2824760:C:TG364D0.997
9:2831319:C:GR181P0.997
9:2831332:G:CH177D0.997
9:2834033:T:AR146S0.997
9:2834033:T:GR146S0.997
9:2834047:A:GW142R0.997
9:2834047:A:TW142R0.997
9:2820049:T:CD413G0.996
9:2820050:C:GD413H0.996
9:2824719:T:CK378E0.996
9:2829838:G:TA263D0.996
9:2830983:A:TV219D0.996
9:2831009:A:CS210R0.996

dbSNP variants (sampled 300 via entrez): RS1000041588 (9:2840647 C>T), RS1000057532 (9:2835835 T>C), RS1000071404 (9:2804641 A>G), RS1000100594 (9:2837806 T>C), RS1000134480 (9:2824130 C>T), RS1000166631 (9:2839149 C>T), RS1000222758 (9:2842197 G>A), RS1000295043 (9:2814893 G>A,C), RS1000302035 (9:2810153 C>G), RS1000364099 (9:2820026 T>A,G), RS1000388103 (9:2815173 T>A), RS1000429986 (9:2835608 G>A), RS1000467686 (9:2823047 T>C), RS1000474777 (9:2837505 C>A,T), RS1000540782 (9:2822864 T>A,C)

Disease associations

OMIM: gene MIM:609960 | disease phenotypes:

GenCC curated gene-disease

Mondo (0):

Orphanet (0):

HPO phenotypes

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

GWAS associations

9 associations (top):

StudyTraitp-value
GCST001762_367Obesity-related traits9.000000e-06
GCST001762_499Obesity-related traits1.000000e-06
GCST002039_5Blood trace element (Se levels)3.000000e-06
GCST002187_8Systolic blood pressure in sickle cell anemia8.000000e-06
GCST005839_13Depression2.000000e-08
GCST006041_17Major depressive disorder4.000000e-08
GCST008161_71Waist circumference adjusted for body mass index7.000000e-06
GCST009391_1188Metabolite levels4.000000e-06
GCST011703_91Smoking initiation2.000000e-08

EFO canonical traits (6, from GWAS)

EFO IDTrait name
EFO:0006336diastolic blood pressure
EFO:0004344birth weight
EFO:0006335systolic blood pressure
EFO:0007789BMI-adjusted waist circumference
EFO:0021575adipic acid measurement
EFO:0005670smoking initiation

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
Tretinoindecreases expression4
bisphenol Aaffects cotreatment, affects methylation, decreases expression3
Benzo(a)pyrenedecreases expression, affects methylation3
Cyclosporineincreases expression2
aristolochic acid Idecreases expression1
bisphenol Faffects cotreatment, increases expression1
dicrotophosdecreases expression1
2,4,6-tribromophenoldecreases expression1
triphenyl phosphateaffects expression1
deoxynivalenolincreases expression1
decabromobiphenyl etherdecreases expression1
sodium arsenitedecreases expression1
tetrabromobisphenol Adecreases expression1
nickel sulfatedecreases expression1
perfluoro-n-nonanoic acidincreases expression1
2,2’,4,4’-tetrabromodiphenyl etherdecreases expression1
pentabrominated diphenyl ether 100decreases expression1
hexabrominated diphenyl ether 153decreases expression1
Resveratrolaffects cotreatment, increases expression1
Fulvestrantaffects cotreatment, affects methylation1
Dichlorodiphenyl Dichloroethyleneincreases expression1
Dexamethasoneaffects cotreatment, increases expression1
Diethylstilbestrolincreases expression1
Dimethyl Sulfoxideaffects expression1
Doxorubicindecreases expression1
Estradiolincreases expression1
Hydrogen Peroxideaffects expression1
Indomethacinaffects cotreatment, increases expression1
Nickelincreases expression1
Plant Extractsaffects cotreatment, increases expression1

Cellosaurus cell lines

2 cell lines: 2 cancer cell line

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

CellosaurusNameCategorySex
CVCL_ZD04Mel 05.18Cancer cell lineMale
CVCL_ZD05Mel 06.07Cancer 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.