MRPL2

gene
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Also known as MRP-L14RPML14CGI-22uL2m

Summary

MRPL2 (mitochondrial ribosomal protein L2, HGNC:14056) is a protein-coding gene on chromosome 6p21.1, encoding Large ribosomal subunit protein uL2m (Q5T653). It is a selective cancer dependency (DepMap: 41.2% of cell lines).

Mammalian mitochondrial ribosomal proteins are encoded by nuclear genes and help in protein synthesis within the mitochondrion. Mitochondrial ribosomes (mitoribosomes) consist of a small 28S subunit and a large 39S subunit. They have an estimated 75% protein to rRNA composition compared to prokaryotic ribosomes, where this ratio is reversed. Another difference between mammalian mitoribosomes and prokaryotic ribosomes is that the latter contain a 5S rRNA. Among different species, the proteins comprising the mitoribosome differ greatly in sequence, and sometimes in biochemical properties, which prevents easy recognition by sequence homology. This gene encodes a 39S subunit protein that belongs to the EcoL2 ribosomal protein family. A pseudogene corresponding to this gene is found on chromosome 12q. Alternative splicing results in multiple transcript variants encoding distinct isoforms.

Source: NCBI Gene 51069 — RefSeq curated summary.

At a glance

  • Clinical variants (ClinVar): 57 total
  • Cancer dependency (DepMap): dependent in 41.2% of screened cell lines
  • MANE Select transcript: NM_015950

Identifiers

Gene identifiers

FieldValue
HGNC IDHGNC:14056
Approved symbolMRPL2
Namemitochondrial ribosomal protein L2
Location6p21.1
Locus typegene with protein product
StatusApproved
AliasesMRP-L14, RPML14, CGI-22, uL2m
Ensembl geneENSG00000112651
Ensembl biotypeprotein_coding
OMIM611822
Entrez51069

Gene structure

Transcript identifiers

Ensembl transcripts: 15 — 11 protein_coding, 4 retained_intron

ENST00000230413, ENST00000388752, ENST00000468957, ENST00000470667, ENST00000480286, ENST00000485654, ENST00000487429, ENST00000489623, ENST00000491898, ENST00000927866, ENST00000927867, ENST00000927868, ENST00000927869, ENST00000927870, ENST00000927871

RefSeq mRNA: 2 — MANE Select: NM_015950 NM_001300848, NM_015950

CCDS: CCDS34454, CCDS75458

Canonical transcript exons

ENST00000388752 — 7 exons

ExonStartEnd
ENSE000007517704305589743056007
ENSE000011380704305402943054486
ENSE000018238664305928643059434
ENSE000034920054305554543055618
ENSE000035243244305630743056445
ENSE000035246664305806543058233
ENSE000035668674305608143056196

Expression profiles

Bgee: expression breadth ubiquitous, 263 present calls, max score 96.92.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 52.6137 / max 413.8488, expressed in 1824 samples.

FANTOM5 promoters (2 alternative TSS)

Promoter IDTPM avgSamples expressed
7366451.39211824
736651.2216752

Top tissues by expression

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

TissueAnatomy IDExpression scoreQuality
hindlimb stylopod muscleUBERON:000425296.92gold quality
mucosa of transverse colonUBERON:000499196.27gold quality
right lobe of liverUBERON:000111495.89gold quality
gastrocnemiusUBERON:000138895.83gold quality
right atrium auricular regionUBERON:000663195.65gold quality
muscle of legUBERON:000138395.52gold quality
apex of heartUBERON:000209895.17gold quality
metanephros cortexUBERON:001053394.81gold quality
transverse colonUBERON:000115794.80gold quality
left adrenal glandUBERON:000123494.74gold quality
adenohypophysisUBERON:000219694.73gold quality
left adrenal gland cortexUBERON:003582594.69gold quality
right adrenal glandUBERON:000123394.63gold quality
cardiac atriumUBERON:000208194.62gold quality
heart left ventricleUBERON:000208494.61gold quality
body of pancreasUBERON:000115094.50gold quality
right lobe of thyroid glandUBERON:000111994.37gold quality
left lobe of thyroid glandUBERON:000112094.36gold quality
right adrenal gland cortexUBERON:003582794.25gold quality
cardiac ventricleUBERON:000208294.17gold quality
lower esophagus muscularis layerUBERON:003583394.13gold quality
lower esophagusUBERON:001347394.12gold quality
muscle layer of sigmoid colonUBERON:003580594.11gold quality
esophagogastric junction muscularis propriaUBERON:003584194.11gold quality
body of stomachUBERON:000116193.78gold quality
olfactory segment of nasal mucosaUBERON:000538693.73gold quality
adrenal cortexUBERON:000123593.65gold quality
endocervixUBERON:000045893.63gold quality
tibial arteryUBERON:000761093.61gold quality
popliteal arteryUBERON:000225093.60gold quality

Single-cell (SCXA)

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

ExperimentMarker?Max mean expression
E-ANND-3yes7.18

Regulation

Is transcription factor: no

miRNA regulators (miRDB)

23 targeting MRPL2, 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-3613-3P100.0076.367965
HSA-MIR-196A-1-3P99.9972.152772
HSA-MIR-3617-3P99.9867.86918
HSA-MIR-3065-5P99.9771.563281
HSA-MIR-3682-5P99.9367.971163
HSA-MIR-6721-5P99.9368.922981
HSA-MIR-3529-3P99.9073.553045
HSA-MIR-472999.6972.184233
HSA-MIR-885-5P99.5968.59879
HSA-MIR-486-5P99.5170.39707
HSA-MIR-7109-5P99.1866.131057
HSA-MIR-328-5P99.0864.651000
HSA-MIR-140-3P99.0467.691324
HSA-MIR-194-5P99.0169.651465
HSA-MIR-6885-5P98.7164.33902
HSA-MIR-6887-5P98.5668.491295
HSA-MIR-6795-5P98.5268.511277
HSA-MIR-3074-3P97.8367.26922
HSA-MIR-192-3P97.5267.661001
HSA-MIR-6892-5P97.2768.60847
HSA-MIR-1915-5P95.2565.78571
HSA-MIR-4707-5P90.9565.69110
HSA-MIR-598-3P89.2567.61112

Functional genomics

DepMap (CRISPR cell-line fitness): dependent in 41.2% of screened cell lines.

Cross-species orthologs

5 orthologs

OrganismSymbolGene ID
danio_reriomrpl2ENSDARG00000062568
mus_musculusMrpl2ENSMUSG00000002767
rattus_norvegicusMrpl2ENSRNOG00000018057
drosophila_melanogastermRpL2FBGN0036135
caenorhabditis_elegansWBGENE00018932

Paralogs (1): RPL8 (ENSG00000161016)

Protein

Protein identifiers

Large ribosomal subunit protein uL2mQ5T653 (reviewed: Q5T653)

Alternative names: 39S ribosomal protein L2, mitochondrial

All UniProt accessions (5): C9IY40, C9J5E0, Q5T653, C9JZG7, C9JZW2

UniProt curated annotations — full annotation on UniProt →

Subunit / interactions. Component of the mitochondrial large ribosomal subunit (mt-LSU). Mature mammalian 55S mitochondrial ribosomes consist of a small (28S) and a large (39S) subunit. The 28S small subunit contains a 12S ribosomal RNA (12S mt-rRNA) and 30 different proteins. The 39S large subunit contains a 16S rRNA (16S mt-rRNA), a copy of mitochondrial valine transfer RNA (mt-tRNA(Val)), which plays an integral structural role, and 52 different proteins.

Subcellular location. Mitochondrion.

Similarity. Belongs to the universal ribosomal protein uL2 family.

RefSeq proteins (2): NP_001287777, NP_057034* (*=MANE)

Domains & families (InterPro)

IDNameType
IPR002171Ribosomal_uL2Family
IPR008991Translation_prot_SH3-like_sfHomologous_superfamily
IPR012340NA-bd_OB-foldHomologous_superfamily
IPR014722Rib_uL2_dom2Homologous_superfamily
IPR022666Ribosomal_uL2_RNA-bd_domDomain
IPR022669Ribosomal_uL2_CDomain

Pfam: PF00181, PF03947

UniProt features (32 total): strand 18, helix 5, turn 2, sequence conflict 2, transit peptide 1, chain 1, region of interest 1, compositionally biased region 1, sequence variant 1

Structure

Experimental structures (PDB)

86 structures, top 30 by resolution.

PDBMethodResolution (Å)
7OF0ELECTRON MICROSCOPY2.2
7QI4ELECTRON MICROSCOPY2.21
8RRIELECTRON MICROSCOPY2.4
8QU5ELECTRON MICROSCOPY2.42
9OLFELECTRON MICROSCOPY2.46
7OF7ELECTRON MICROSCOPY2.5
7PO4ELECTRON MICROSCOPY2.56
6ZM6ELECTRON MICROSCOPY2.59
7O9MELECTRON MICROSCOPY2.6
7OF6ELECTRON MICROSCOPY2.6
9CN3ELECTRON MICROSCOPY2.62
7QI5ELECTRON MICROSCOPY2.63
7OF2ELECTRON MICROSCOPY2.7
7OF3ELECTRON MICROSCOPY2.7
7OF4ELECTRON MICROSCOPY2.7
8QU1ELECTRON MICROSCOPY2.74
9PR4ELECTRON MICROSCOPY2.77
9PRAELECTRON MICROSCOPY2.83
8ANYELECTRON MICROSCOPY2.85
6ZM5ELECTRON MICROSCOPY2.89
7QH7ELECTRON MICROSCOPY2.89
7ODRELECTRON MICROSCOPY2.9
7OF5ELECTRON MICROSCOPY2.9
8K2AELECTRON MICROSCOPY2.9
8OITELECTRON MICROSCOPY2.9
9PGLELECTRON MICROSCOPY2.9
9PGFELECTRON MICROSCOPY2.93
6VMIELECTRON MICROSCOPY2.96
6VLZELECTRON MICROSCOPY2.97
7QI6ELECTRON MICROSCOPY2.98

Predicted structure (AlphaFold)

ModelpLDDTFraction very-high
AF-Q5T653-F186.030.75

Function

Pathways and Gene Ontology

Reactome pathways

7 pathways

IDPathway
R-HSA-5368286Mitochondrial translation initiation
R-HSA-5389840Mitochondrial translation elongation
R-HSA-5419276Mitochondrial translation termination
R-HSA-9937383Mitochondrial ribosome-associated quality control
R-HSA-392499Metabolism of proteins
R-HSA-5368287Mitochondrial translation
R-HSA-72766Translation

MSigDB gene sets: 122 (showing top): GOBP_MITOCHONDRIAL_TRANSLATION, TGACCTY_ERR1_Q2, GGGTGGRR_PAX4_03, COUP_01, GOBP_TRANSLATION, GOMF_STRUCTURAL_CONSTITUENT_OF_RIBOSOME, HNF4_DR1_Q3, GOCC_MITOCHONDRIAL_ENVELOPE, CTGYNNCTYTAA_UNKNOWN, HAMAI_APOPTOSIS_VIA_TRAIL_DN, GOCC_LARGE_RIBOSOMAL_SUBUNIT, GOCC_RIBOSOME, GOCC_ORGANELLAR_RIBOSOME, GOCC_MITOCHONDRIAL_MATRIX, GOCC_ORGANELLE_INNER_MEMBRANE

GO Biological Process (2): mitochondrial translation (GO:0032543), translation (GO:0006412)

GO Molecular Function (2): RNA binding (GO:0003723), structural constituent of ribosome (GO:0003735)

GO Cellular Component (6): nucleoplasm (GO:0005654), mitochondrion (GO:0005739), mitochondrial inner membrane (GO:0005743), mitochondrial large ribosomal subunit (GO:0005762), ribosome (GO:0005840), ribonucleoprotein complex (GO:1990904)

Reactome top-level categories

Rollup of top-3 pathways:

CategoryPathways
Mitochondrial translation4
Translation1
Metabolism of proteins1

GO top-level categories

Rollup of top GO terms by namespace:

CategoryTerms
mitochondrion1
translation1
mitochondrial gene expression1
peptidyltransferase activity1
translational initiation1
translational elongation1
translational termination1
macromolecule biosynthetic process1
protein metabolic process1
protein biosynthetic process1
nucleic acid binding1
structural molecule activity1
ribosome1
nuclear lumen1
cellular anatomical structure1
cytoplasm1
intracellular membrane-bounded organelle1
organelle inner membrane1
mitochondrial membrane1
organellar large ribosomal subunit1
mitochondrial ribosome1
mitochondrial protein-containing complex1
intracellular membraneless organelle1
protein-containing complex1

Protein interactions and networks

STRING

4596 interactions, top by confidence (×1000):

Protein AProtein BPartner UniProtScore
MRPL2MRPL16Q9NX20662
MRPL2MRPS18AQ9NVS2646
MRPL2MRPL1Q9BYD6593
MRPL2IMP4Q96G21576
MRPL2MRPS12O15235565
MRPL2MRPL9Q9BYD2534
MRPL2MRPL17Q9NRX2527
MRPL2MRPL21Q7Z2W9496
MRPL2MRPS16Q9Y3D3489
MRPL2MRPL11Q9Y3B7478
MRPL2PEX5LQ8IYB4473
MRPL2MRPS6P82932472
MRPL2MRPS2Q9Y399470
MRPL2NSA2O95478465
MRPL2MRPS10P82664453

IntAct

148 interactions, top by confidence:

ABTypeScore
YBX1HNRNPRpsi-mi:“MI:0914”(association)0.770
EZH2EPOPpsi-mi:“MI:0914”(association)0.730
MALSU1NDUFAB1psi-mi:“MI:0914”(association)0.710
MRPS30GTPBP10psi-mi:“MI:0914”(association)0.640
NPKPNA6psi-mi:“MI:0914”(association)0.550
MRPL50GTPBP10psi-mi:“MI:0914”(association)0.530
MRPL10ZZEF1psi-mi:“MI:0914”(association)0.530
MRPL42GATCpsi-mi:“MI:0914”(association)0.530
MRPL28MRPL3psi-mi:“MI:0914”(association)0.530
NSA2TYW5psi-mi:“MI:0914”(association)0.530
MRPL2GTPBP10psi-mi:“MI:0914”(association)0.530
MRPL13GTPBP10psi-mi:“MI:0914”(association)0.530
MRPL18GTPBP10psi-mi:“MI:0914”(association)0.530
MRPL16MRPL2psi-mi:“MI:0914”(association)0.530
MRPL23BCKDHApsi-mi:“MI:0914”(association)0.530
MRPL27MRPL33psi-mi:“MI:0914”(association)0.530
NDUFAB1MIEF1psi-mi:“MI:0915”(physical association)0.490
CMTM4MRPL2psi-mi:“MI:0915”(physical association)0.370
TRIM63MRPL2psi-mi:“MI:0915”(physical association)0.370
MRPL2SMURF2psi-mi:“MI:0915”(physical association)0.370
MASTLMED26psi-mi:“MI:0914”(association)0.350
Brwd3WDR91psi-mi:“MI:0914”(association)0.350
NcstnDERL1psi-mi:“MI:0914”(association)0.350
MRPL50MRPL43psi-mi:“MI:0914”(association)0.350

BioGRID (301): MRPL2 (Affinity Capture-MS), MRPL2 (Affinity Capture-MS), MRPL2 (Affinity Capture-MS), MRPL2 (Affinity Capture-MS), MRPL2 (Affinity Capture-MS), MRPL2 (Affinity Capture-MS), MRPL2 (Affinity Capture-MS), GFM2 (Co-fractionation), GTPBP4 (Co-fractionation), MRPL13 (Co-fractionation), MRPL17 (Co-fractionation), MRPL2 (Co-fractionation), MRPL2 (Co-fractionation), MRPL2 (Co-fractionation), MRPL2 (Co-fractionation)

ESM2 similar proteins: A0AJY7, B1VKG7, O60783, P56556, P82915, P82920, P82933, Q02366, Q0MQA3, Q0MQA4, Q0MQA5, Q0VFH6, Q13084, Q19V89, Q1JQ99, Q29IK4, Q2HJJ1, Q2TA12, Q2TBR2, Q3T040, Q3T0J3, Q498T4, Q58DQ5, Q58DV5, Q5I0K8, Q5T653, Q5U4Z8, Q5ZMU0, Q6B860, Q6DDY9, Q6P1L8, Q71YD4, Q7M0E7, Q7SF85, Q7ZUH5, Q80X85, Q8MFA1, Q8Y620, Q92AC7, Q9BYC9

Diamond homologs: A0ALW5, A0L5X6, A0LIJ3, A0R8I3, A1ALU4, A1VEB3, A4IJJ2, A4XLS7, A5GAW8, A5IM86, A5USI6, A6KYJ2, A6LEI8, A6LLL6, A7GK23, A7HBM1, A7HWR4, A7IFY4, A7NR60, A8F4R4, A8ZV60, A9B415, A9BHA2, A9WH69, B0K5P6, B0KCK3, B1LBN7, B1YGV3, B2RLY9, B3E7T8, B3EUM0, B3PK40, B4S094, B4S5M4, B4UBA2, B5EFQ3, B5Y985, B5YDU6, B6IRQ9, B7HQU7

SIGNOR signaling

1 interactions.

AEffectBMechanism
MRPL2“form complex”“39S mitochondrial large ribosomal subunit”binding

Enriched among interaction partners

Reactome pathways and GO biological processes over-represented among this gene’s 139 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
Mitochondrial ribosome-associated quality control3844.9×6e-53
Mitochondrial translation3343.7×2e-45
Mitochondrial translation initiation3542.7×6e-48
Mitochondrial translation elongation3542.7×6e-48
Mitochondrial translation termination3537.0×2e-45
Translation3520.9×8e-36
Peptide chain elongation911.0×5e-06
Viral mRNA Translation911.0×5e-06

GO biological processes:

GO termPartnersFoldFDR
mitochondrial translation3751.8×1e-52
mitochondrial large ribosomal subunit assembly540.0×2e-05
translation2621.6×3e-25
cytoplasmic translation913.4×4e-06
negative regulation of translation69.5×5e-03

Disease & clinical

Clinical variants and AI predictions

ClinVar

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

ClassificationCount (floor)
Pathogenic0
Likely pathogenic0
Uncertain significance44
Likely benign2
Benign0

Top pathogenic / likely-pathogenic (0)

SpliceAI

997 predictions. Top by Δscore:

VariantEffectΔscore
6:43055893:ATAC:Adonor_loss1.0000
6:43055894:TACC:Tdonor_loss1.0000
6:43055895:A:ATdonor_loss1.0000
6:43055896:C:CTdonor_loss1.0000
6:43055934:A:ACdonor_gain1.0000
6:43055935:C:CCdonor_gain1.0000
6:43056005:CAA:Cacceptor_gain1.0000
6:43056008:C:CCacceptor_gain1.0000
6:43056075:TCTCA:Tdonor_loss1.0000
6:43056076:CTCA:Cdonor_loss1.0000
6:43056077:TCA:Tdonor_loss1.0000
6:43056078:CA:Cdonor_loss1.0000
6:43056079:ACC:Adonor_loss1.0000
6:43056080:C:CAdonor_loss1.0000
6:43056195:ACC:Aacceptor_loss1.0000
6:43058061:CCAC:Cdonor_loss1.0000
6:43058063:A:Tdonor_loss1.0000
6:43058064:C:CTdonor_loss1.0000
6:43058089:T:TAdonor_gain1.0000
6:43058090:C:Adonor_gain1.0000
6:43058095:T:Cdonor_gain1.0000
6:43058234:C:CCacceptor_gain1.0000
6:43054486:CCT:Cacceptor_loss0.9900
6:43054488:T:Cacceptor_loss0.9900
6:43055614:CGTCC:Cacceptor_gain0.9900
6:43055619:C:CCacceptor_gain0.9900
6:43055938:T:Adonor_gain0.9900
6:43056003:AGCAA:Aacceptor_gain0.9900
6:43056004:GCAA:Gacceptor_gain0.9900
6:43056005:CAAC:Cacceptor_gain0.9900

AlphaMissense

1955 scored. Top likely-pathogenic:

VariantProtein changeam_pathogenicity
6:43056153:A:GW150R0.993
6:43056153:A:TW150R0.993
6:43056183:C:GA140P0.991
6:43054467:G:TA242E0.990
6:43055609:C:TG214D0.989
6:43055897:C:AG211W0.989
6:43056155:C:GR149P0.989
6:43056182:G:TA140D0.989
6:43055573:A:TI226N0.988
6:43056307:C:GR135T0.986
6:43054472:G:CC240W0.985
6:43055567:A:GL228P0.985
6:43055597:C:GR218P0.985
6:43056307:C:AR135M0.985
6:43055618:C:AG211V0.984
6:43055950:A:TI193N0.984
6:43056196:C:AR135S0.984
6:43056196:C:GR135S0.984
6:43056176:A:TV142D0.983
6:43056191:G:TA137E0.983
6:43054392:C:AG267V0.982
6:43054458:C:AG245V0.982
6:43054458:C:TG245D0.982
6:43054459:C:GG245R0.982
6:43055610:C:GG214R0.982
6:43055977:A:TL184H0.982
6:43055899:G:TA210E0.981
6:43056185:A:TI139K0.981
6:43056334:A:TV126D0.981
6:43054473:C:TC240Y0.980

dbSNP variants (sampled 300 via entrez): RS1000434098 (6:43053884 G>A,C), RS1000465372 (6:43054079 A>C), RS1000810369 (6:43060798 G>A), RS1001162643 (6:43055172 G>A), RS1001426967 (6:43054951 G>A), RS1002704714 (6:43059099 T>A), RS1003236245 (6:43058211 T>C), RS1003297582 (6:43057267 A>C,G), RS1003370186 (6:43055679 C>A,G,T), RS1004142730 (6:43060884 GAC>G), RS1004824855 (6:43057478 G>A), RS1005100018 (6:43057063 T>C), RS1005596433 (6:43054520 T>C), RS1005767494 (6:43059768 C>T), RS1007047532 (6:43060909 C>G)

Disease associations

OMIM: gene MIM:611822 | disease phenotypes:

GenCC curated gene-disease

Mondo (0):

Orphanet (0):

HPO phenotypes

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

GWAS associations

0 associations (top):

Drugs & pharmacology

Drug and pharmacology data

Is drug target: no

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

CTD chemical–gene interactions

26 total (human), top 26 by PubMed support.

ChemicalActions (top 5)PubMed papers
sodium arsenitedecreases expression, affects cotreatment, increases abundance, increases expression2
aristolochic acid Iincreases expression1
TAK-243increases sumoylation1
methylmercuric chloridedecreases expression1
triphenyl phosphateaffects expression1
arseniteaffects binding, increases reaction1
manganese chlorideaffects cotreatment, increases abundance, increases expression1
perfluorooctane sulfonic acidincreases expression1
CGP 52608affects binding, increases reaction1
ICG 001decreases expression1
4-(4-((5-(4,5-dimethyl-2-nitrophenyl)-2-furanyl)methylene)-4,5-dihydro-3-methyl-5-oxo-1H-pyrazol-1-yl)benzoic aciddecreases expression1
Temozolomideincreases expression1
Air Pollutantsdecreases expression, increases abundance1
Arsenicaffects cotreatment, increases abundance, increases expression1
Atrazineincreases expression1
Benzo(a)pyrenedecreases expression1
Ivermectindecreases expression1
Manganeseaffects cotreatment, increases abundance, increases expression1
Ribonucleotidesaffects binding1
Seleniumaffects cotreatment, increases expression1
Smokedecreases expression, increases abundance1
Valproic Acidincreases expression1
Vitamin Eaffects cotreatment, increases expression1
Gold Compoundsdecreases methylation1
Acrylamidedecreases expression1
tert-Butylhydroperoxideincreases expression1

Cellosaurus cell lines

1 cell lines: 1 transformed cell line

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

CellosaurusNameCategorySex
CVCL_B3B9Abcam HEK293T MRPL2 KOTransformed cell lineFemale

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.