STRBP

gene
On this page

Also known as FLJ11307SPNR

Summary

STRBP (spermatid perinuclear RNA binding protein, HGNC:16462) is a protein-coding gene on chromosome 9q33.3, encoding Spermatid perinuclear RNA-binding protein (Q96SI9). Involved in spermatogenesis and sperm function.

Enables RNA binding activity. Predicted to be involved in cell differentiation and spermatogenesis. Predicted to act upstream of or within cell motility; mechanosensory behavior; and spermatid development. Located in nucleus.

Source: NCBI Gene 55342 — RefSeq curated summary.

At a glance

  • GWAS associations: 8
  • Clinical variants (ClinVar): 73 total
  • MANE Select transcript: NM_018387

Identifiers

Gene identifiers

FieldValue
HGNC IDHGNC:16462
Approved symbolSTRBP
Namespermatid perinuclear RNA binding protein
Location9q33.3
Locus typegene with protein product
StatusApproved
AliasesFLJ11307, SPNR
Ensembl geneENSG00000165209
Ensembl biotypeprotein_coding
OMIM611138
Entrez55342

Gene structure

Transcript identifiers

Ensembl transcripts: 29 — 23 protein_coding, 4 nonsense_mediated_decay, 1 retained_intron, 1 protein_coding_CDS_not_defined

ENST00000348403, ENST00000360998, ENST00000407982, ENST00000447404, ENST00000471564, ENST00000478973, ENST00000479114, ENST00000492885, ENST00000496533, ENST00000530364, ENST00000878273, ENST00000878274, ENST00000878275, ENST00000878276, ENST00000878277, ENST00000878278, ENST00000878279, ENST00000878280, ENST00000932652, ENST00000932653, ENST00000932654, ENST00000932655, ENST00000932656, ENST00000932657, ENST00000932658, ENST00000932659, ENST00000932660, ENST00000970350, ENST00000970351

RefSeq mRNA: 5 — MANE Select: NM_018387 NM_001171137, NM_001376106, NM_001376107, NM_001376109, NM_018387

CCDS: CCDS55337, CCDS6851

Canonical transcript exons

ENST00000348403 — 19 exons

ExonStartEnd
ENSE00001287138123173677123173842
ENSE00001327887123236830123236966
ENSE00001728022123121617123125673
ENSE00003473955123147778123147870
ENSE00003481471123158332123158429
ENSE00003485892123184132123184298
ENSE00003522375123160367123160462
ENSE00003539553123159096123159207
ENSE00003547336123136381123136515
ENSE00003560758123139529123139687
ENSE00003564023123146855123147054
ENSE00003576993123132845123132968
ENSE00003591136123128214123128258
ENSE00003610091123158012123158123
ENSE00003620842123136041123136181
ENSE00003672110123160977123161068
ENSE00003689174123169902123170046
ENSE00003787305123179007123179227
ENSE00003890679123268436123268586

Expression profiles

Bgee: expression breadth ubiquitous, 253 present calls, max score 98.47.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 24.7166 / max 602.8518, expressed in 1685 samples.

FANTOM5 promoters (4 alternative TSS)

Promoter IDTPM avgSamples expressed
10240823.37131682
1024071.1526403
1024040.184573
1024030.00812

Top tissues by expression

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

TissueAnatomy IDExpression scoreQuality
secondary oocyteCL:000065598.47gold quality
primary visual cortexUBERON:000243698.17gold quality
occipital lobeUBERON:000202197.68gold quality
oocyteCL:000002397.14gold quality
cortical plateUBERON:000534396.73gold quality
lateral nuclear group of thalamusUBERON:000273696.53gold quality
Brodmann (1909) area 23UBERON:001355496.31gold quality
spermCL:000001995.99gold quality
ganglionic eminenceUBERON:000402395.19gold quality
endothelial cellCL:000011595.07gold quality
oviduct epitheliumUBERON:000480494.68gold quality
parietal lobeUBERON:000187294.66gold quality
postcentral gyrusUBERON:000258194.60gold quality
epithelium of nasopharynxUBERON:000195194.56gold quality
parotid glandUBERON:000183194.48gold quality
bronchial epithelial cellCL:000232894.39gold quality
right uterine tubeUBERON:000130294.10gold quality
superior frontal gyrusUBERON:000266194.09gold quality
bronchusUBERON:000218594.04gold quality
ileal mucosaUBERON:000033193.78gold quality
left testisUBERON:000453393.38gold quality
olfactory segment of nasal mucosaUBERON:000538693.27gold quality
right testisUBERON:000453493.19gold quality
mucosa of paranasal sinusUBERON:000503092.88gold quality
middle temporal gyrusUBERON:000277192.68gold quality
prefrontal cortexUBERON:000045192.64gold quality
lower esophagus mucosaUBERON:003583492.59gold quality
Brodmann (1909) area 9UBERON:001354092.49gold quality
dorsolateral prefrontal cortexUBERON:000983492.48gold quality
testisUBERON:000047392.35gold quality

Single-cell (SCXA)

Detected in 2 experiment(s), a significant marker in 2.

ExperimentMarker?Max mean expression
E-CURD-112yes37.49
E-ANND-3yes15.07

Regulation

Is transcription factor: no

miRNA regulators (miRDB)

260 targeting STRBP, 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-8485100.0077.574731
HSA-MIR-4668-3P100.0068.742635
HSA-MIR-6867-5P100.0082.213464
HSA-MIR-4776-3P100.0068.731340
HSA-MIR-3613-3P100.0076.367965
HSA-MIR-3646100.0073.565283
HSA-MIR-6833-3P100.0070.633197
HSA-MIR-4768-5P100.0069.492861
HSA-MIR-513A-5P100.0069.772465
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-5692A100.0074.406850
HSA-MIR-4795-3P100.0074.624024
HSA-MIR-1277-5P100.0073.955056
HSA-MIR-450099.9972.722367
HSA-MIR-196A-1-3P99.9972.152772
HSA-MIR-513B-5P99.9969.962150
HSA-MIR-511-3P99.9968.851467
HSA-MIR-366299.9973.825684
HSA-MIR-520D-5P99.9873.344883
HSA-MIR-524-5P99.9873.434882
HSA-MIR-1213699.9872.815713
HSA-MIR-477599.9875.006394
HSA-LET-7A-5P99.9872.291790
HSA-LET-7B-5P99.9872.311790
HSA-LET-7C-5P99.9872.291790
HSA-LET-7E-5P99.9872.291790
HSA-LET-7F-5P99.9872.561784

Literature-anchored findings (GeneRIF, showing 3)

  • Low STRBP mRNA is associated with cryptorchidism and Down’s syndrome. (PMID:22391137)
  • Spermatid perinuclear RNA-binding protein promotes UBR5-mediated proteolysis of Dicer to accelerate triple-negative breast cancer progression. (PMID:38280476)
  • CircSTRBP contributes to H2O2-induced lens epithelium cell dysfunction through increasing NOX4 mRNA stability by recruiting IGF2BP1. (PMID:38340945)

Cross-species orthologs

8 orthologs

OrganismSymbolGene ID
danio_reriostrbpENSDARG00000021455
mus_musculusStrbpENSMUSG00000026915
rattus_norvegicusStrbpENSRNOG00000010150
drosophila_melanogasterloqsFBGN0032515
drosophila_melanogasterCG12493FBGN0035571
drosophila_melanogasterblanksFBGN0035608
drosophila_melanogasterZn72DFBGN0263603
caenorhabditis_eleganszfr-1WBGENE00022388

Paralogs (14): STAU2 (ENSG00000040341), ZFR (ENSG00000056097), ADAT1 (ENSG00000065457), ZFR2 (ENSG00000105278), STAU1 (ENSG00000124214), ILF3 (ENSG00000129351), TARBP2 (ENSG00000139546), ADAD2 (ENSG00000140955), ILF2 (ENSG00000143621), ADAR (ENSG00000160710), ADAD1 (ENSG00000164113), PRKRA (ENSG00000180228), ADARB2 (ENSG00000185736), ADARB1 (ENSG00000197381)

Protein

Protein identifiers

Spermatid perinuclear RNA-binding proteinQ96SI9 (reviewed: Q96SI9)

All UniProt accessions (8): A0A0S2Z5K8, E9PKQ0, Q96SI9, H0YC91, H0YD00, H0YD70, Q5JPA5, V9HWK4

UniProt curated annotations — full annotation on UniProt →

Function. Involved in spermatogenesis and sperm function. Plays a role in regulation of cell growth. Binds to double-stranded DNA and RNA. Binds most efficiently to poly(I:C) RNA than to poly(dI:dC) DNA. Binds also to single-stranded poly(G) RNA. Binds non-specifically to the mRNA PRM1 3’-UTR and adenovirus VA RNA.

Subunit / interactions. Interacts with EIF2AK2. Associates with microtubules; it is unsure whether such interaction is direct or indirect.

Subcellular location. Cytoplasm.

Isoforms (2)

UniProt IDNamesCanonical?
Q96SI9-11yes
Q96SI9-22

RefSeq proteins (5): NP_001164608, NP_001363035, NP_001363036, NP_001363038, NP_060857* (*=MANE)

Domains & families (InterPro)

IDNameType
IPR006561DZF_domDomain
IPR014720dsRBD_domDomain
IPR043519NT_sfHomologous_superfamily
IPR044472STRBP_DSRM_1Domain
IPR049401DZF_dom_NDomain
IPR049402DZF_dom_CDomain

Pfam: PF00035, PF07528, PF20965

UniProt features (23 total): strand 5, domain 3, region of interest 3, compositionally biased region 3, modified residue 2, sequence conflict 2, helix 2, chain 1, splice variant 1, sequence variant 1

Structure

Experimental structures (PDB)

1 structures.

PDBMethodResolution (Å)
2DMYSOLUTION NMR

Predicted structure (AlphaFold)

ModelpLDDTFraction very-high
AF-Q96SI9-F173.420.51

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 (2): 612, 617

Function

Pathways and Gene Ontology

Reactome pathways

0 pathways

MSigDB gene sets: 251 (showing top): GOBP_BEHAVIOR, STARK_PREFRONTAL_CORTEX_22Q11_DELETION_DN, GOZGIT_ESR1_TARGETS_DN, GAUSSMANN_MLL_AF4_FUSION_TARGETS_A_UP, ATGCAGT_MIR217, GOBP_MALE_GAMETE_GENERATION, MARTINEZ_RB1_TARGETS_UP, MUELLER_PLURINET, MILI_PSEUDOPODIA_HAPTOTAXIS_UP, LIAO_METASTASIS, GOBP_CELLULAR_PROCESS_INVOLVED_IN_REPRODUCTION_IN_MULTICELLULAR_ORGANISM, GARCIA_TARGETS_OF_FLI1_AND_DAX1_DN, GOBP_RESPONSE_TO_ABIOTIC_STIMULUS, RYTTCCTG_ETS2_B, GOBP_DEVELOPMENTAL_PROCESS_INVOLVED_IN_REPRODUCTION

GO Biological Process (5): spermatid development (GO:0007286), mechanosensory behavior (GO:0007638), cell motility (GO:0048870), spermatogenesis (GO:0007283), cell differentiation (GO:0030154)

GO Molecular Function (7): DNA binding (GO:0003677), RNA binding (GO:0003723), double-stranded RNA binding (GO:0003725), single-stranded RNA binding (GO:0003727), microtubule binding (GO:0008017), protein binding (GO:0005515), tubulin binding (GO:0015631)

GO Cellular Component (4): manchette (GO:0002177), nucleus (GO:0005634), cytoplasm (GO:0005737), microtubule cytoskeleton (GO:0015630)

GO top-level categories

Rollup of top GO terms by namespace:

CategoryTerms
nucleic acid binding2
RNA binding2
cellular anatomical structure2
germ cell development1
spermatid differentiation1
behavior1
response to mechanical stimulus1
cellular process1
developmental process involved in reproduction1
male gamete generation1
cellular developmental process1
tubulin binding1
binding1
cytoskeletal protein binding1
microtubule cytoskeleton1
intracellular membrane-bounded organelle1
intracellular anatomical structure1
cytoskeleton1

Protein interactions and networks

STRING

1230 interactions, top by confidence (×1000):

Protein AProtein BPartner UniProtScore
STRBPEIF2AK2P19525759
STRBPDDX17Q92841696
STRBPPOLDIP3Q9BY77685
STRBPPIF1Q9H611667
STRBPPRM1P04553563
STRBPU2AF2P26368535
STRBPA0A087WZY1A0A087WZY1516
STRBPPRH1P02810507
STRBPPOLA1P09884452
STRBPZNF398Q8TD17450
STRBPGPR21Q99679431
STRBPDYNLT1P63172429
STRBPTHAP1Q9NVV9428
STRBPNUP62P37198427
STRBPUNC79Q9P2D8426
STRBPSPAG1Q07617426

IntAct

167 interactions, top by confidence:

ABTypeScore
HNRNPCKPNA3psi-mi:“MI:0914”(association)0.850
THAP1STRBPpsi-mi:“MI:0915”(physical association)0.780
STRBPTHAP1psi-mi:“MI:0915”(physical association)0.780
CFTRESYT2psi-mi:“MI:0914”(association)0.710
TARBP2STRBPpsi-mi:“MI:0915”(physical association)0.670
STRBPRBPMSpsi-mi:“MI:0915”(physical association)0.670
STRBPTARBP2psi-mi:“MI:0915”(physical association)0.670
PRKRASTRBPpsi-mi:“MI:0915”(physical association)0.670
RBPMSSTRBPpsi-mi:“MI:0915”(physical association)0.670
IGF2BP1IGF2BP3psi-mi:“MI:0914”(association)0.640
RAC1COX6Cpsi-mi:“MI:0914”(association)0.640
ASF1BHAT1psi-mi:“MI:0914”(association)0.640
CFTRHAX1psi-mi:“MI:0914”(association)0.610
DAB1STRBPpsi-mi:“MI:0915”(physical association)0.560
STRBPDAB1psi-mi:“MI:0915”(physical association)0.560
STRBPRBPMSpsi-mi:“MI:0915”(physical association)0.560
STRBPRBFOX1psi-mi:“MI:0915”(physical association)0.560
STRBPZMAT4psi-mi:“MI:0915”(physical association)0.560

BioGRID (294): STRBP (Two-hybrid), STRBP (Two-hybrid), STRBP (Two-hybrid), STRBP (Two-hybrid), STRBP (Affinity Capture-MS), STRBP (Affinity Capture-MS), STRBP (Affinity Capture-MS), STRBP (Affinity Capture-MS), STRBP (Affinity Capture-MS), STRBP (Affinity Capture-MS), STRBP (Affinity Capture-MS), STRBP (Two-hybrid), STRBP (Two-hybrid), STRBP (Affinity Capture-RNA), STRBP (Affinity Capture-MS)

ESM2 similar proteins: A0A8C0TYJ0, A2AWA9, A2VDZ9, A5GFS8, O70133, O76031, O88506, O95292, O95747, Q08E27, Q0VCY1, Q12959, Q13330, Q15691, Q16514, Q3T174, Q3ZBD9, Q5PYH5, Q5R495, Q5R601, Q5R6Y5, Q5R7N3, Q5R7Z5, Q5U2U0, Q5ZHN3, Q5ZIL4, Q61166, Q61187, Q62599, Q62696, Q66HR2, Q68FK8, Q6AY57, Q6IRE4, Q6ZVM7, Q7ZWU5, Q80W47, Q811D0, Q863I2, Q8K4B0

Diamond homologs: O88532, Q08E27, Q0VD35, Q12906, Q562A2, Q5R6Y5, Q5REX3, Q5U231, Q5ZIL4, Q6DCD0, Q6DD04, Q6GL57, Q6GPM1, Q6NXA4, Q6PCR6, Q7TP98, Q91550, Q91WM1, Q96KR1, Q96SI9, Q9H898, Q9JIL3, Q9JKU6, Q9UPR6, Q9Z1X4, A2RFW8, A8AWC2, B5XKB7, B8GAM6, C0MCR4, P0DF14, P0DF15, P51400, P55265, P55266, P66670, P66672, P78563, P97473, P97616

SIGNOR signaling

0 interactions.

Enriched among interaction partners

Reactome pathways and GO biological processes over-represented among this gene’s 162 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
PKR-mediated signaling79.3×9e-04
mRNA Polyadenylation97.5×6e-04
Processing of Capped Intron-Containing Pre-mRNA97.0×8e-04
mRNA Splicing - Major Pathway126.2×1e-04
Major pathway of rRNA processing in the nucleolus and cytosol105.8×9e-04
Dengue Virus-Host Interactions135.6×1e-04

GO biological processes:

GO termPartnersFoldFDR
positive regulation of cytoplasmic translation534.9×8e-05
positive regulation of viral genome replication520.5×5e-04
negative regulation of translation1013.8×3e-06
RNA processing812.3×8e-05
ribosomal small subunit biogenesis711.2×4e-04
positive regulation of translation711.2×4e-04
regulation of alternative mRNA splicing, via spliceosome610.3×3e-03
translation85.8×7e-03

Disease & clinical

Clinical variants and AI predictions

ClinVar

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

ClassificationCount (floor)
Pathogenic0
Likely pathogenic0
Uncertain significance49
Likely benign1
Benign0

Top pathogenic / likely-pathogenic (0)

SpliceAI

3813 predictions. Top by Δscore:

VariantEffectΔscore
9:123128729:T:Adonor_gain1.0000
9:123136035:TCATA:Tdonor_loss1.0000
9:123136036:CATAC:Cdonor_loss1.0000
9:123136038:TACCT:Tdonor_loss1.0000
9:123136039:ACC:Adonor_loss1.0000
9:123136040:C:CGdonor_loss1.0000
9:123136182:CTAC:Cacceptor_loss1.0000
9:123136183:T:Cacceptor_loss1.0000
9:123136376:CACA:Cdonor_loss1.0000
9:123136377:ACACC:Adonor_loss1.0000
9:123136378:CACCT:Cdonor_loss1.0000
9:123136379:A:Cdonor_loss1.0000
9:123136379:ACCT:Adonor_gain1.0000
9:123136380:C:Gdonor_loss1.0000
9:123136380:CCTC:Cdonor_gain1.0000
9:123136511:CTTAC:Cacceptor_gain1.0000
9:123136514:ACC:Aacceptor_loss1.0000
9:123136516:C:CGacceptor_loss1.0000
9:123146850:TGTA:Tdonor_loss1.0000
9:123146852:TAC:Tdonor_loss1.0000
9:123146853:A:AGdonor_loss1.0000
9:123147055:C:CCacceptor_gain1.0000
9:123147773:CA:Cdonor_gain1.0000
9:123147773:CACA:Cdonor_gain1.0000
9:123147776:A:ACdonor_gain1.0000
9:123147777:C:CTdonor_gain1.0000
9:123147777:CTTTT:Cdonor_gain1.0000
9:123147781:T:TAdonor_gain1.0000
9:123147786:A:ACdonor_gain1.0000
9:123147787:A:Cdonor_gain1.0000

AlphaMissense

4381 scored. Top likely-pathogenic:

VariantProtein changeam_pathogenicity
9:123136093:A:GL574P1.000
9:123136102:A:CL571W1.000
9:123136102:A:GL571S1.000
9:123136105:G:TA570D1.000
9:123136106:C:GA570P1.000
9:123136114:G:AA567V1.000
9:123136114:G:TA567D1.000
9:123136115:C:GA567P1.000
9:123136115:C:TA567T1.000
9:123136117:G:TA566E1.000
9:123136118:C:GA566P1.000
9:123136123:G:TA564E1.000
9:123136124:C:GA564P1.000
9:123136125:C:AK563N1.000
9:123136125:C:GK563N1.000
9:123136126:T:AK563M1.000
9:123136126:T:GK563T1.000
9:123136127:T:CK563E1.000
9:123136127:T:GK563Q1.000
9:123136129:G:AA562V1.000
9:123136129:G:TA562E1.000
9:123136130:C:GA562P1.000
9:123136130:C:TA562T1.000
9:123136134:T:AK560N1.000
9:123136134:T:GK560N1.000
9:123136135:T:AK560I1.000
9:123136136:T:CK560E1.000
9:123136136:T:GK560Q1.000
9:123136137:C:AK559N1.000
9:123136137:C:GK559N1.000

dbSNP variants (sampled 300 via entrez): RS1000007828 (9:123162001 C>T), RS1000011260 (9:123235886 T>C), RS1000018521 (9:123128650 T>A,C), RS1000029776 (9:123208110 CTG>C), RS1000035919 (9:123146094 A>G), RS1000054217 (9:123114715 T>C), RS1000057392 (9:123177990 A>T), RS1000087821 (9:123256923 T>C), RS1000089836 (9:123147377 C>A), RS1000136686 (9:123230862 C>T), RS1000158278 (9:123203396 C>T), RS1000186378 (9:123187173 C>G), RS1000205511 (9:123256704 C>G), RS1000220308 (9:123205910 C>T), RS1000234452 (9:123213088 A>G)

Disease associations

OMIM: gene MIM:611138 | disease phenotypes:

GenCC curated gene-disease

Mondo (0):

Orphanet (0):

HPO phenotypes

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

GWAS associations

8 associations (top):

StudyTraitp-value
GCST005146_13Birth weight1.000000e-12
GCST005951_64Body mass index2.000000e-10
GCST008053_108Height2.000000e-11
GCST008053_186Height4.000000e-07
GCST008839_63Height1.000000e-12
GCST009616_3HDL cholesterol levels x thiazide or thiazide-like diuretics use interaction7.000000e-07
GCST010703_296Brain morphology (MOSTest)2.000000e-14
GCST90002388_227Lymphocyte count2.000000e-10

EFO canonical traits (5, from GWAS)

EFO IDTrait name
EFO:0004344birth weight
EFO:0004340body mass index
EFO:0004612high density lipoprotein cholesterol measurement
EFO:0004346neuroimaging measurement
EFO:0004587lymphocyte count

Drugs & pharmacology

Drug and pharmacology data

Is drug target: no

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

PharmGKB clinical annotations

1 annotations.

VariantTypeLevelDrugsPhenotypes
rs9032Efficacy3methylphenidateAttention Deficit Disorder with Hyperactivity

CTD chemical–gene interactions

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

ChemicalActions (top 5)PubMed papers
Tetrachlorodibenzodioxindecreases expression, increases expression3
Phenylmercuric Acetateaffects cotreatment, decreases expression2
Cadmium Chloridedecreases expression, increases expression2
aristolochic acid Idecreases expression1
GSK-J4increases expression1
dicrotophosdecreases expression1
urushiolincreases expression1
methylmercuric chloridedecreases expression1
triphenyl phosphateaffects expression1
bisphenol Adecreases expression1
deoxynivalenolincreases expression1
cobaltous chloridedecreases expression1
benzo(e)pyrenedecreases methylation1
di-n-butylphosphoric acidaffects expression1
CGP 52608increases reaction, affects binding1
4-(5-benzo(1,3)dioxol-5-yl-4-pyridin-2-yl-1H-imidazol-2-yl)benzamideaffects cotreatment, decreases expression1
ICG 001increases expression1
abrinedecreases expression1
dorsomorphinaffects cotreatment, decreases expression1
Arsenic Trioxideincreases expression1
Cidofovirdecreases expression1
Acetaminophendecreases expression1
Air Pollutantsincreases abundance, increases expression1
Atrazineincreases expression1
Benzenedecreases expression1
Caffeinedecreases expression1
Diazinonincreases methylation1
Doxorubicindecreases expression1
Enzyme Inhibitorsdecreases activity, increases O-linked glycosylation1
Estradiolaffects cotreatment, increases 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.