WDR75

gene
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Also known as FLJ12519NET16UTP17

Summary

WDR75 (WD repeat domain 75, HGNC:25725) is a protein-coding gene on chromosome 2q32.2, encoding WD repeat-containing protein 75 (Q8IWA0). Ribosome biogenesis factor. It is a common-essential gene (DepMap: required in 99.6% of cancer cell lines).

Enables RNA binding activity. Involved in positive regulation of rRNA processing; positive regulation of transcription by RNA polymerase I; and ribosomal small subunit biogenesis. Located in nucleolus. Part of small-subunit processome.

Source: NCBI Gene 84128 — RefSeq curated summary.

At a glance

  • GWAS associations: 10
  • Clinical variants (ClinVar): 106 total
  • Cancer dependency (DepMap): dependent in 99.6% of screened cell lines (common-essential)
  • MANE Select transcript: NM_032168

Identifiers

Gene identifiers

FieldValue
HGNC IDHGNC:25725
Approved symbolWDR75
NameWD repeat domain 75
Location2q32.2
Locus typegene with protein product
StatusApproved
AliasesFLJ12519, NET16, UTP17
Ensembl geneENSG00000115368
Ensembl biotypeprotein_coding
OMIM620341
Entrez84128

Gene structure

Transcript identifiers

Ensembl transcripts: 13 — 8 protein_coding, 3 retained_intron, 2 nonsense_mediated_decay

ENST00000314761, ENST00000427960, ENST00000436347, ENST00000472286, ENST00000475596, ENST00000498365, ENST00000631047, ENST00000873640, ENST00000922924, ENST00000922925, ENST00000922926, ENST00000922927, ENST00000922928

RefSeq mRNA: 2 — MANE Select: NM_032168 NM_001303096, NM_032168

CCDS: CCDS2298

Canonical transcript exons

ENST00000314761 — 21 exons

ExonStartEnd
ENSE00001070666189441474189441578
ENSE00001233905189475213189475552
ENSE00003462757189463694189463753
ENSE00003494394189466425189466582
ENSE00003494695189469344189469439
ENSE00003495544189458753189458872
ENSE00003504762189470819189470878
ENSE00003507228189474717189474808
ENSE00003507351189468475189468569
ENSE00003513124189463846189463961
ENSE00003536714189467468189467648
ENSE00003537595189474186189474332
ENSE00003539395189462484189462642
ENSE00003577705189450903189450968
ENSE00003589568189457311189457381
ENSE00003605154189448379189448508
ENSE00003606347189465079189465254
ENSE00003639176189459336189459424
ENSE00003655735189451805189451895
ENSE00003662349189455320189455444
ENSE00003677252189470076189470245

Expression profiles

Bgee: expression breadth ubiquitous, 260 present calls, max score 93.58.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 29.9946 / max 429.4280, expressed in 1803 samples.

FANTOM5 promoters (2 alternative TSS)

Promoter IDTPM avgSamples expressed
2425126.74681797
242503.24781379

Top tissues by expression

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

TissueAnatomy IDExpression scoreQuality
upper arm skinUBERON:000426393.58gold quality
cardiac muscle of right atriumUBERON:000337993.47gold quality
left ventricle myocardiumUBERON:000656693.30gold quality
kidney epitheliumUBERON:000481993.02gold quality
myocardiumUBERON:000234992.51gold quality
cartilage tissueUBERON:000241892.45gold quality
smooth muscle tissueUBERON:000113592.31gold quality
primordial germ cell in gonadCL:0000670 ∩ UBERON:000099192.27gold quality
calcaneal tendonUBERON:000370192.18gold quality
body of uterusUBERON:000985392.07gold quality
left ovaryUBERON:000211992.04gold quality
right ovaryUBERON:000211891.98gold quality
oocyteCL:000002391.72gold quality
endometriumUBERON:000129591.43gold quality
ovaryUBERON:000099291.32gold quality
mucosa of stomachUBERON:000119991.28gold quality
endocervixUBERON:000045891.22gold quality
uterusUBERON:000099591.12gold quality
lymph nodeUBERON:000002991.00gold quality
superficial temporal arteryUBERON:000161491.00gold quality
rectumUBERON:000105290.99gold quality
left uterine tubeUBERON:000130390.94gold quality
corpus callosumUBERON:000233690.77gold quality
mucosa of paranasal sinusUBERON:000503090.73gold quality
ectocervixUBERON:001224990.64gold quality
uterine cervixUBERON:000000290.62gold quality
left adrenal gland cortexUBERON:003582590.53gold quality
right uterine tubeUBERON:000130290.51gold quality
body of pancreasUBERON:000115090.47gold quality
adrenal tissueUBERON:001830390.43gold quality

Single-cell (SCXA)

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

ExperimentMarker?Max mean expression
E-ANND-3yes4.97

Regulation

Is transcription factor: no

miRNA regulators (miRDB)

9 targeting WDR75, 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-806899.9873.852376
HSA-MIR-314399.9371.963104
HSA-MIR-10527-5P99.9172.283754
HSA-MIR-548E-5P99.8972.734486
HSA-MIR-205399.5769.151635
HSA-MIR-432899.5771.064094
HSA-MIR-374A-3P98.8767.821531
HSA-MIR-477398.3567.301710
HSA-MIR-217-3P95.6768.421000

Functional genomics

DepMap (CRISPR cell-line fitness): dependent in 99.6% of screened cell lines, common-essential.

Literature-anchored findings (GeneRIF, showing 1)

  • RNA-interference screen for p53 regulators unveils a role of WDR75 in ribosome biogenesis. (PMID:34611297)

Cross-species orthologs

3 orthologs

OrganismSymbolGene ID
danio_reriowdr75ENSDARG00000040730
mus_musculusWdr75ENSMUSG00000025995
rattus_norvegicusWdr75ENSRNOG00000003825

Paralogs (26): PAFAH1B1 (ENSG00000007168), SNRNP40 (ENSG00000060688), WDR62 (ENSG00000075702), WDR7 (ENSG00000091157), TBL2 (ENSG00000106638), PAK1IP1 (ENSG00000111845), DCAF4 (ENSG00000119599), DAW1 (ENSG00000123977), TEP1 (ENSG00000129566), AHI1 (ENSG00000135541), WDR38 (ENSG00000136918), MAPKBP1 (ENSG00000137802), POC1B (ENSG00000139323), NEDD1 (ENSG00000139350), COP1 (ENSG00000143207), WDR17 (ENSG00000150627), WDR43 (ENSG00000163811), POC1A (ENSG00000164087), WDR88 (ENSG00000166359), WDR81 (ENSG00000167716), DCAF4L2 (ENSG00000176566), DCAF4L1 (ENSG00000182308), WDR27 (ENSG00000184465), NWD1 (ENSG00000188039), WDR5 (ENSG00000196363), WDR5B (ENSG00000196981)

Protein

Protein identifiers

WD repeat-containing protein 75Q8IWA0 (reviewed: Q8IWA0)

Alternative names: U3 small nucleolar RNA-associated protein 17 homolog

All UniProt accessions (3): Q8IWA0, F8WC81, F8WE20

UniProt curated annotations — full annotation on UniProt →

Function. Ribosome biogenesis factor. Part of the small subunit (SSU) processome, first precursor of the small eukaryotic ribosomal subunit. During the assembly of the SSU processome in the nucleolus, many ribosome biogenesis factors, an RNA chaperone and ribosomal proteins associate with the nascent pre-rRNA and work in concert to generate RNA folding, modifications, rearrangements and cleavage as well as targeted degradation of pre-ribosomal RNA by the RNA exosome. Involved in nucleolar processing of pre-18S ribosomal RNA. Required for optimal pre-ribosomal RNA transcription by RNA polymerase I.

Subunit / interactions. Component of the proposed t-UTP subcomplex of the ribosomal small subunit (SSU) processome. SSU processome is composed of more than 70 proteins and the RNA chaperone small nucleolar RNA (snoRNA) U3.

Subcellular location. Nucleus. Nucleolus.

RefSeq proteins (2): NP_001290025, NP_115544* (*=MANE)

Domains & families (InterPro)

IDNameType
IPR001680WD40_rptRepeat
IPR011047Quinoprotein_ADH-like_sfHomologous_superfamily
IPR015943WD40/YVTN_repeat-like_dom_sfHomologous_superfamily
IPR053826WDR75Family
IPR057644Beta-prop_WDR75_2ndDomain

Pfam: PF23769, PF23869

UniProt features (33 total): repeat 13, modified residue 7, sequence conflict 7, cross-link 3, chain 1, region of interest 1, compositionally biased region 1

Structure

Experimental structures (PDB)

3 structures.

PDBMethodResolution (Å)
7MQAELECTRON MICROSCOPY2.7
7MQ8ELECTRON MICROSCOPY3.6
7MQ9ELECTRON MICROSCOPY3.87

Predicted structure (AlphaFold)

ModelpLDDTFraction very-high
AF-Q8IWA0-F180.430.49

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 (10): 466, 664, 672, 779, 782, 796, 811, 123, 427, 676

Function

Pathways and Gene Ontology

Reactome pathways

2 pathways

IDPathway
R-HSA-6790901rRNA modification in the nucleus and cytosol
R-HSA-6791226Major pathway of rRNA processing in the nucleolus and cytosol

MSigDB gene sets: 155 (showing top): GSE45365_NK_CELL_VS_BCELL_UP, MODULE_97, GOBP_RIBOSOME_BIOGENESIS, RODRIGUES_THYROID_CARCINOMA_ANAPLASTIC_UP, RODRIGUES_THYROID_CARCINOMA_POORLY_DIFFERENTIATED_UP, GOBP_MATURATION_OF_SSU_RRNA, GRAESSMANN_APOPTOSIS_BY_SERUM_DEPRIVATION_UP, GRAESSMANN_APOPTOSIS_BY_DOXORUBICIN_DN, GRAESSMANN_RESPONSE_TO_MC_AND_DOXORUBICIN_DN, MODULE_182, WEI_MYCN_TARGETS_WITH_E_BOX, MARTINEZ_RB1_TARGETS_DN, MILI_PSEUDOPODIA_HAPTOTAXIS_UP, GOBP_RIBOSOMAL_SMALL_SUBUNIT_BIOGENESIS, DODD_NASOPHARYNGEAL_CARCINOMA_UP

GO Biological Process (5): rRNA processing (GO:0006364), ribosomal small subunit biogenesis (GO:0042274), positive regulation of transcription by RNA polymerase I (GO:0045943), positive regulation of rRNA processing (GO:2000234), ribosome biogenesis (GO:0042254)

GO Molecular Function (2): RNA binding (GO:0003723), protein binding (GO:0005515)

GO Cellular Component (4): nucleoplasm (GO:0005654), nucleolus (GO:0005730), small-subunit processome (GO:0032040), nucleus (GO:0005634)

Reactome top-level categories

Rollup of top-1 pathways:

CategoryPathways
rRNA processing in the nucleus and cytosol2

GO top-level categories

Rollup of top GO terms by namespace:

CategoryTerms
ribosome biogenesis2
ribonucleoprotein complex biogenesis2
nuclear lumen2
RNA processing1
rRNA metabolic process1
regulation of transcription by RNA polymerase I1
transcription by RNA polymerase I1
positive regulation of DNA-templated transcription1
rRNA processing1
positive regulation of RNA metabolic process1
regulation of rRNA processing1
nucleic acid binding1
binding1
cellular anatomical structure1
intracellular membraneless organelle1
nucleolus1
preribosome1
t-UTP complex1
nuclear protein-containing complex1
intracellular membrane-bounded organelle1

Protein interactions and networks

STRING

2353 interactions, top by confidence (×1000):

Protein AProtein BPartner UniProtScore
WDR75UTP15Q8TED0941
WDR75WDR43Q15061935
WDR75UTP4Q969X6919
WDR75HEATR1Q9H583914
WDR75MPHOSPH10O00566703
WDR75WDR36Q8NI36665
WDR75UTP18Q9Y5J1646
WDR75UTP20O75691635
WDR75WDR46O15213625
WDR75NOP58Q9Y2X3589
WDR75SNU13P55769579
WDR75BMS1Q14692579
WDR75UTP6Q9NYH9578
WDR75PWP2Q15269576
WDR75TBL3Q12788560

IntAct

97 interactions, top by confidence:

ABTypeScore
C1DZFC3H1psi-mi:“MI:0914”(association)0.640
CACNB1CACNB3psi-mi:“MI:0914”(association)0.640
NIPAL1ESYT2psi-mi:“MI:0914”(association)0.640
EXOSC5ZFC3H1psi-mi:“MI:0914”(association)0.640
EXOSC3MTREXpsi-mi:“MI:0914”(association)0.640
NOL7WDR43psi-mi:“MI:0914”(association)0.640
PWP2FBLpsi-mi:“MI:0914”(association)0.610
PLEKWDR75psi-mi:“MI:0915”(physical association)0.590
ASB7POLR3Apsi-mi:“MI:0914”(association)0.530
SERPINA12TSPAN6psi-mi:“MI:0914”(association)0.530
NOL11WDR43psi-mi:“MI:0914”(association)0.530
WDR75WDR43psi-mi:“MI:0914”(association)0.530
FBLZNF316psi-mi:“MI:0914”(association)0.530
UTP4ANKRD28psi-mi:“MI:0914”(association)0.530
RBM4NVLpsi-mi:“MI:0914”(association)0.530
WDR43WDR75psi-mi:“MI:0914”(association)0.530
NOL7IPO5psi-mi:“MI:0914”(association)0.530
MPHOSPH6ZFC3H1psi-mi:“MI:0914”(association)0.530
Smad3psi-mi:“MI:0914”(association)0.350
CEP170P1PCYT1Apsi-mi:“MI:0914”(association)0.350
ATP6AP2TMUB1psi-mi:“MI:0914”(association)0.350
Srp72psi-mi:“MI:0914”(association)0.350
KIF21Bpsi-mi:“MI:0914”(association)0.350
Setdb1INPP5Bpsi-mi:“MI:0914”(association)0.350
EXOSC5PAPD5psi-mi:“MI:0914”(association)0.350

BioGRID (144): WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS), WDR75 (Affinity Capture-MS)

ESM2 similar proteins: A0A0G2JEB6, A0JM56, B0DOB4, B0FXQ5, B1ANS9, B4F7L9, B4GQJ7, B5DHW4, B7FF06, B7FF07, B7FF08, B7FF09, B7FF12, C5IAW9, F1LW30, F1P4W9, O08747, O95185, P0DM40, Q008S8, Q18264, Q32NR9, Q3V0B4, Q402B2, Q4G0P3, Q5R4M2, Q5T0N1, Q5XI14, Q6AXU1, Q6DCF6, Q6NRS1, Q6P2C0, Q6P5D8, Q6UXZ4, Q6ZTR5, Q6ZU64, Q761X5, Q7T2Z5, Q80W93, Q86YR7

Diamond homologs: A4RDD7, A6ZPA6, A7EF03, A7THX0, B3LJT5, B4HSL3, B4QHG6, B8P4B0, B8PD53, B9WD30, C4YPI7, C5DF48, C5DY07, C5FP68, C5MJE8, C7GWC1, C8ZH19, D3TLL6, D4AM37, D4D8P3, O00628, P39946, P97865, Q54WA3, Q5A7Q3, Q5A7Q6, Q6CJ50, Q6CU55, Q6DFC6, Q6FT96, Q6FWT9, Q759U7, Q7KNS3, Q7RXH4, Q8IWA0, Q8R537, Q8RXA7, Q9H2Y7, A0JMQ0, A2QP30

SIGNOR signaling

1 interactions.

AEffectBMechanism
WDR75“form complex”“UTP-A complex”binding

Enriched among interaction partners

Reactome pathways and GO biological processes over-represented among this gene’s 129 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
Nuclear RNA decay725.4×1e-06
rRNA modification in the nucleus and cytosol919.8×1e-07
Major pathway of rRNA processing in the nucleolus and cytosol1712.3×9e-12
mRNA Splicing911.6×5e-06
Processing of Capped Intron-Containing Pre-mRNA109.7×5e-06
mRNA Splicing - Major Pathway127.7×4e-06
Metabolism of RNA146.9×1e-06

GO biological processes:

GO termPartnersFoldFDR
maturation of SSU-rRNA532.5×6e-05
RNA catabolic process519.3×6e-04
ribosomal small subunit biogenesis917.4×7e-07
rRNA processing1113.2×4e-07
RNA processing713.0×1e-04
regulation of alternative mRNA splicing, via spliceosome612.4×8e-04
RNA splicing118.2×2e-05
mRNA splicing, via spliceosome86.2×2e-03

Disease & clinical

Clinical variants and AI predictions

ClinVar

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

ClassificationCount (floor)
Pathogenic0
Likely pathogenic0
Uncertain significance76
Likely benign0
Benign0

Top pathogenic / likely-pathogenic (0)

SpliceAI

3245 predictions. Top by Δscore:

VariantEffectΔscore
2:189441553:A:Tdonor_gain1.0000
2:189441571:GATTC:Gdonor_gain1.0000
2:189441575:C:CGdonor_gain1.0000
2:189441579:G:GGdonor_gain1.0000
2:189448378:GGT:Gacceptor_gain1.0000
2:189448378:GGTAT:Gacceptor_gain1.0000
2:189450901:A:AGacceptor_gain1.0000
2:189450902:G:GGacceptor_gain1.0000
2:189450902:GCT:Gacceptor_gain1.0000
2:189450902:GCTGT:Gacceptor_gain1.0000
2:189455318:A:AGacceptor_gain1.0000
2:189455319:G:GGacceptor_gain1.0000
2:189455319:GA:Gacceptor_gain1.0000
2:189455319:GAT:Gacceptor_gain1.0000
2:189455319:GATA:Gacceptor_gain1.0000
2:189455319:GATAT:Gacceptor_gain1.0000
2:189455441:CGAGG:Cdonor_loss1.0000
2:189455442:GAG:Gdonor_gain1.0000
2:189455442:GAGG:Gdonor_loss1.0000
2:189455445:GT:Gdonor_loss1.0000
2:189455446:T:Adonor_loss1.0000
2:189458905:T:Gdonor_gain1.0000
2:189460374:G:GTdonor_gain1.0000
2:189460374:G:Tdonor_gain1.0000
2:189462347:A:Gdonor_gain1.0000
2:189462445:A:AGacceptor_gain1.0000
2:189462446:A:AGacceptor_gain1.0000
2:189462447:A:Gacceptor_gain1.0000
2:189462448:A:Gacceptor_gain1.0000
2:189462449:C:Gacceptor_gain1.0000

AlphaMissense

5496 scored. Top likely-pathogenic:

VariantProtein changeam_pathogenicity
2:189450939:T:AW85R0.999
2:189450939:T:CW85R0.999
2:189462498:A:CS265R0.999
2:189462500:T:AS265R0.999
2:189462500:T:GS265R0.999
2:189466552:T:AW473R0.999
2:189466552:T:CW473R0.999
2:189467482:T:AW488R0.999
2:189467482:T:CW488R0.999
2:189458857:G:TG225V0.998
2:189458871:T:AW230R0.998
2:189458871:T:CW230R0.998
2:189462528:T:AW275R0.998
2:189462528:T:CW275R0.998
2:189463897:T:AV350D0.998
2:189463921:T:CL358P0.998
2:189467590:T:AW524R0.998
2:189467590:T:CW524R0.998
2:189458839:C:AA219E0.997
2:189458857:G:AG225D0.997
2:189458866:G:CR228P0.997
2:189459382:T:AW246R0.997
2:189459382:T:CW246R0.997
2:189465170:T:CL402P0.997
2:189466520:T:AV462D0.997
2:189470204:T:AW650R0.997
2:189470204:T:CW650R0.997
2:189458845:G:AG221D0.996
2:189462520:T:CL272P0.996
2:189465229:T:AW422R0.996

dbSNP variants (sampled 300 via entrez): RS1000100795 (2:189441546 C>A), RS1000265209 (2:189472693 G>A), RS1000342115 (2:189448036 T>C), RS1000472594 (2:189459359 A>C), RS1000551485 (2:189440011 G>A,C), RS1000714829 (2:189453921 G>A), RS1000991207 (2:189440830 C>G), RS1001024489 (2:189471151 C>G), RS1001193860 (2:189457445 T>C), RS1001269404 (2:189470969 T>C), RS1001308476 (2:189457679 G>A), RS1001339920 (2:189441093 C>A,G,T), RS1001422374 (2:189464193 T>C), RS1001472604 (2:189458079 A>T), RS1001524740 (2:189457904 C>T)

Disease associations

OMIM: gene MIM:620341 | disease phenotypes:

GenCC curated gene-disease

Mondo (0):

Orphanet (0):

HPO phenotypes

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

GWAS associations

10 associations (top):

StudyTraitp-value
GCST002677_1Iron status biomarkers (ferritin levels)8.000000e-19
GCST002678_1Iron status biomarkers (transferrin levels)1.000000e-11
GCST004571_22Iron status biomarkers (total iron binding capacity)8.000000e-07
GCST004572_3Iron status biomarkers (transferrin saturation)8.000000e-07
GCST010698_39Subcortical volume (min-P)9.000000e-26
GCST010699_36Brain morphology (min-P)9.000000e-09
GCST010700_26Cortical thickness (MOSTest)9.000000e-52
GCST010701_48Cortical surface area (MOSTest)2.000000e-18
GCST010702_50Subcortical volume (MOSTest)8.000000e-10
GCST010703_340Brain morphology (MOSTest)3.000000e-10

EFO canonical traits (6, from GWAS)

EFO IDTrait name
EFO:0004459ferritin measurement
EFO:0004461iron biomarker measurement
EFO:0006341transferrin measurement
EFO:0006334total iron binding capacity
EFO:0004346neuroimaging measurement
EFO:0004840cortical thickness

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
sodium arseniteincreases expression, decreases expression, increases abundance3
Valproic Aciddecreases expression2
Cadmium Chloridedecreases expression2
FR900359affects phosphorylation1
dicrotophosdecreases expression1
alpha phellandrenedecreases expression1
triphenyl phosphateaffects expression1
alpha-pineneaffects cotreatment, increases expression, increases abundance1
propionaldehydeincreases methylation1
deoxynivalenolincreases expression1
nonanalincreases methylation1
n-hexanalincreases methylation1
beta-lapachoneincreases expression1
arsenitedecreases reaction, affects binding1
butyraldehydeincreases methylation1
caprylic aldehydeincreases methylation1
nivalenolincreases expression1
methacrylaldehydeaffects cotreatment, increases expression, increases abundance1
pentanalincreases methylation1
heptanalincreases methylation1
di-n-butylphosphoric acidaffects expression1
CGP 52608affects binding, increases reaction1
jinfukangdecreases expression1
(+)-JQ1 compoundincreases expression1
Resveratrolaffects cotreatment, increases expression1
Acroleinaffects cotreatment, increases expression, increases abundance1
Air Pollutantsaffects cotreatment, increases abundance, increases expression1
Arsenicincreases abundance, increases expression1
Doxorubicindecreases expression1
Estradiolincreases 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.