ZNF564
gene geneOn this page
Also known as MGC26914
Summary
ZNF564 (zinc finger protein 564, HGNC:31106) is a protein-coding gene on chromosome 19p13.2, encoding Zinc finger protein 564 (Q8TBZ8). May be involved in transcriptional regulation.
Predicted to enable DNA-binding transcription factor activity and RNA polymerase II cis-regulatory region sequence-specific DNA binding activity. Predicted to be involved in regulation of transcription by RNA polymerase II. Predicted to be located in nucleus.
Source: NCBI Gene 163050 — RefSeq curated summary.
At a glance
- Clinical variants (ClinVar): 76 total
- MANE Select transcript:
NM_144976
Identifiers
Gene identifiers
| Field | Value |
|---|---|
| HGNC ID | HGNC:31106 |
| Approved symbol | ZNF564 |
| Name | zinc finger protein 564 |
| Location | 19p13.2 |
| Locus type | gene with protein product |
| Status | Approved |
| Aliases | MGC26914 |
| Ensembl gene | ENSG00000249709 |
| Ensembl biotype | protein_coding |
| Entrez | 163050 |
Gene structure
Transcript identifiers
Ensembl transcripts: 5 — 3 protein_coding, 1 nonsense_mediated_decay, 1 retained_intron
ENST00000339282, ENST00000416136, ENST00000427105, ENST00000465410, ENST00000596193
RefSeq mRNA: 1 — MANE Select: NM_144976
NM_144976
CCDS: CCDS42505
Canonical transcript exons
ENST00000339282 — 4 exons
| Exon | Start | End |
|---|---|---|
| ENSE00001429098 | 12525373 | 12527916 |
| ENSE00001529909 | 12551330 | 12551482 |
| ENSE00003634566 | 12528304 | 12528364 |
| ENSE00003682173 | 12528570 | 12528696 |
Expression profiles
Bgee: expression breadth ubiquitous, 134 present calls, max score 89.90.
FANTOM5 (CAGE): breadth ubiquitous, TPM avg 9.2985 / max 137.5056, expressed in 1748 samples.
FANTOM5 promoters (2 alternative TSS)
| Promoter ID | TPM avg | Samples expressed |
|---|---|---|
| 179314 | 8.5044 | 1733 |
| 179315 | 0.7940 | 505 |
Top tissues by expression
134 total, by Bgee expression score (0-100, higher = more expressed):
| Tissue | Anatomy ID | Expression score | Quality |
|---|---|---|---|
| calcaneal tendon | UBERON:0003701 | 89.90 | gold quality |
| primordial germ cell in gonad | CL:0000670 ∩ UBERON:0000991 | 88.30 | gold quality |
| colonic epithelium | UBERON:0000397 | 85.87 | gold quality |
| corpus callosum | UBERON:0002336 | 85.73 | gold quality |
| sural nerve | UBERON:0015488 | 83.35 | gold quality |
| endometrium | UBERON:0001295 | 82.06 | gold quality |
| mucosa of transverse colon | UBERON:0004991 | 81.55 | gold quality |
| tonsil | UBERON:0002372 | 81.52 | gold quality |
| male germ line stem cell (sensu Vertebrata) in testis | CL:0000089 ∩ UBERON:0000473 | 81.23 | gold quality |
| skeletal muscle tissue | UBERON:0001134 | 80.93 | gold quality |
| urinary bladder | UBERON:0001255 | 80.23 | gold quality |
| muscle tissue | UBERON:0002385 | 79.99 | gold quality |
| ventricular zone | UBERON:0003053 | 79.98 | gold quality |
| leukocyte | CL:0000738 | 79.20 | gold quality |
| rectum | UBERON:0001052 | 79.13 | gold quality |
| monocyte | CL:0000576 | 79.11 | gold quality |
| adrenal tissue | UBERON:0018303 | 79.08 | gold quality |
| bone marrow cell | CL:0002092 | 78.91 | gold quality |
| adult mammalian kidney | UBERON:0000082 | 78.80 | gold quality |
| kidney | UBERON:0002113 | 78.78 | gold quality |
| blood | UBERON:0000178 | 78.48 | gold quality |
| uterine cervix | UBERON:0000002 | 78.36 | gold quality |
| ganglionic eminence | UBERON:0004023 | 78.31 | gold quality |
| lymph node | UBERON:0000029 | 78.25 | gold quality |
| spleen | UBERON:0002106 | 78.16 | gold quality |
| myometrium | UBERON:0001296 | 78.02 | gold quality |
| body of uterus | UBERON:0009853 | 77.93 | gold quality |
| granulocyte | CL:0000094 | 77.85 | gold quality |
| muscle of leg | UBERON:0001383 | 77.66 | gold quality |
| right lobe of thyroid gland | UBERON:0001119 | 77.63 | gold quality |
Single-cell (SCXA)
Detected in 1 experiment(s), a significant marker in 0.
| Experiment | Marker? | Max mean expression |
|---|---|---|
| E-ANND-3 | no | 3.26 |
Regulation
Is transcription factor: no
miRNA regulators (miRDB)
48 targeting ZNF564, top 30 by miRDB confidence (max_score; target_count = how many genes the miRNA targets in total — lower means more specific):
| miRNA | Max score | Avg score | miRNA target_count |
|---|---|---|---|
| HSA-MIR-6870-5P | 99.99 | 68.55 | 2115 |
| HSA-MIR-4531 | 99.99 | 69.70 | 3181 |
| HSA-MIR-4723-5P | 99.97 | 68.70 | 2034 |
| HSA-MIR-5698 | 99.97 | 68.49 | 2029 |
| HSA-MIR-7111-5P | 99.97 | 68.48 | 2062 |
| HSA-MIR-4778-3P | 99.93 | 70.40 | 1818 |
| HSA-MIR-6780A-5P | 99.88 | 66.69 | 2776 |
| HSA-MIR-4447 | 99.85 | 67.81 | 2900 |
| HSA-MIR-4645-3P | 99.76 | 69.33 | 993 |
| HSA-MIR-1255A | 99.74 | 68.09 | 744 |
| HSA-MIR-1255B-5P | 99.74 | 68.16 | 741 |
| HSA-MIR-4446-5P | 99.72 | 69.19 | 2544 |
| HSA-MIR-4524A-3P | 99.72 | 66.85 | 2406 |
| HSA-MIR-33A-3P | 99.70 | 70.27 | 3362 |
| HSA-MIR-6892-3P | 99.68 | 66.40 | 1178 |
| HSA-MIR-452-5P | 99.65 | 69.63 | 1762 |
| HSA-MIR-4676-3P | 99.65 | 69.31 | 1733 |
| HSA-MIR-892C-3P | 99.65 | 69.38 | 1745 |
| HSA-MIR-587 | 99.64 | 70.86 | 2611 |
| HSA-MIR-4472 | 99.56 | 66.08 | 1478 |
| HSA-MIR-190A-5P | 99.54 | 71.45 | 933 |
| HSA-MIR-190B-5P | 99.54 | 71.40 | 925 |
| HSA-MIR-7159-5P | 99.53 | 72.12 | 2472 |
| HSA-MIR-12131 | 99.48 | 68.72 | 1673 |
| HSA-MIR-578 | 99.46 | 68.36 | 1787 |
| HSA-MIR-145-3P | 99.33 | 67.66 | 764 |
| HSA-MIR-1912-3P | 99.32 | 67.40 | 936 |
| HSA-MIR-3978 | 99.24 | 68.39 | 2201 |
| HSA-MIR-4291 | 99.20 | 68.88 | 2969 |
| HSA-MIR-3940-5P | 99.14 | 65.26 | 493 |
Literature-anchored findings (GeneRIF, showing 1)
- Long non-coding RNA ZFAS1 alleviates bupivacaine-induced neurotoxicity by regulating the miR-421/zinc finger protein564 (ZNF564) axis. (PMID:34414857)
Cross-species orthologs
2 orthologs
| Organism | Symbol | Gene ID |
|---|---|---|
| drosophila_melanogaster | CG14442 | FBGN0029893 |
| drosophila_melanogaster | CG14440 | FBGN0029894 |
Paralogs (11): IKZF2 (ENSG00000030419), ZNF821 (ENSG00000102984), ZNF639 (ENSG00000121864), IKZF4 (ENSG00000123411), ZNF382 (ENSG00000161298), IKZF3 (ENSG00000161405), ZNF613 (ENSG00000176024), IKZF1 (ENSG00000185811), ZNF567 (ENSG00000189042), ZNF649 (ENSG00000198093), ZNF350 (ENSG00000256683)
Protein
Protein identifiers
Zinc finger protein 564 — Q8TBZ8 (reviewed: Q8TBZ8)
All UniProt accessions (4): C9JLM4, Q8TBZ8, F8WD42, M0QZN8
UniProt curated annotations — full annotation on UniProt →
Function. May be involved in transcriptional regulation.
Subcellular location. Nucleus.
Similarity. Belongs to the krueppel C2H2-type zinc-finger protein family.
RefSeq proteins (1): NP_659413* (*=MANE)
Domains & families (InterPro)
| ID | Name | Type |
|---|---|---|
| IPR001909 | KRAB | Domain |
| IPR013087 | Znf_C2H2_type | Domain |
| IPR036051 | KRAB_dom_sf | Homologous_superfamily |
| IPR036236 | Znf_C2H2_sf | Homologous_superfamily |
Pfam: PF00096, PF01352, PF13465, PF13912
UniProt features (18 total): zinc finger region 15, chain 1, domain 1, cross-link 1
Structure
Experimental structures (PDB)
0 structures.
Predicted structure (AlphaFold)
| Model | pLDDT | Fraction very-high |
|---|---|---|
| AF-Q8TBZ8-F1 | 77.89 | 0.41 |
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): 438
Function
Pathways and Gene Ontology
Reactome pathways
1 pathways
| ID | Pathway |
|---|---|
| R-HSA-212436 | Generic Transcription Pathway |
MSigDB gene sets: 44 (showing top):
chr19p13, GOMF_SEQUENCE_SPECIFIC_DNA_BINDING, WAMUNYOKOLI_OVARIAN_CANCER_LMP_UP, ZWANG_DOWN_BY_2ND_EGF_PULSE, GOMF_TRANSCRIPTION_REGULATOR_ACTIVITY, ASH1L_TARGET_GENES, CEBPZ_TARGET_GENES, CREB3L4_TARGET_GENES, E2F2_TARGET_GENES, FEV_TARGET_GENES, GREB1_TARGET_GENES, H1_6_TARGET_GENES, PRKDC_TARGET_GENES, SALL4_TARGET_GENES, SOX3_TARGET_GENES
GO Biological Process (2): regulation of transcription by RNA polymerase II (GO:0006357), regulation of DNA-templated transcription (GO:0006355)
GO Molecular Function (6): RNA polymerase II cis-regulatory region sequence-specific DNA binding (GO:0000978), DNA-binding transcription factor activity (GO:0003700), zinc ion binding (GO:0008270), DNA binding (GO:0003677), protein binding (GO:0005515), metal ion binding (GO:0046872)
GO Cellular Component (1): nucleus (GO:0005634)
Reactome top-level categories
Rollup of top-1 pathways:
| Category | Pathways |
|---|---|
| RNA Polymerase II Transcription | 1 |
GO top-level categories
Rollup of top GO terms by namespace:
| Category | Terms |
|---|---|
| regulation of DNA-templated transcription | 2 |
| transcription by RNA polymerase II | 1 |
| DNA-templated transcription | 1 |
| regulation of gene expression | 1 |
| regulation of RNA biosynthetic process | 1 |
| RNA polymerase II transcription regulatory region sequence-specific DNA binding | 1 |
| cis-regulatory region sequence-specific DNA binding | 1 |
| transcription cis-regulatory region binding | 1 |
| transcription regulator activity | 1 |
| transition metal ion binding | 1 |
| nucleic acid binding | 1 |
| binding | 1 |
| cation binding | 1 |
| intracellular membrane-bounded organelle | 1 |
Protein interactions and networks
STRING
402 interactions, top by confidence (×1000):
| Protein A | Protein B | Partner UniProt | Score |
|---|---|---|---|
| ZNF564 | DCAF5 | Q96JK2 | 444 |
| ZNF564 | TRAPPC6B | Q86SZ2 | 434 |
| ZNF564 | YPEL1 | O60688 | 432 |
| ZNF564 | SLC2A4RG | Q9NR83 | 381 |
| ZNF564 | HELB | Q8NG08 | 367 |
| ZNF564 | FOXS1 | O43638 | 360 |
| ZNF564 | FOXJ2 | Q9P0K8 | 317 |
| ZNF564 | SOX14 | O95416 | 307 |
| ZNF564 | HORMAD2 | Q8N7B1 | 291 |
| ZNF564 | TUT1 | Q9H6E5 | 282 |
| ZNF564 | GIGYF2 | Q6Y7W6 | 271 |
| ZNF564 | FCGBP | Q9Y6R7 | 271 |
| ZNF564 | MEF2B | Q02080 | 256 |
| ZNF564 | CKAP5 | Q14008 | 247 |
| ZNF564 | ELK3 | P41970 | 246 |
IntAct
143 interactions, top by confidence:
| A | B | Type | Score |
|---|---|---|---|
| ZNF564 | PPP1R16B | psi-mi:“MI:0915”(physical association) | 0.740 |
| ZNF564 | ALAS1 | psi-mi:“MI:0915”(physical association) | 0.720 |
| TRIM23 | ZNF564 | psi-mi:“MI:0915”(physical association) | 0.720 |
| ZNF564 | KRTAP10-7 | psi-mi:“MI:0915”(physical association) | 0.720 |
| TRAF1 | ZNF564 | psi-mi:“MI:0915”(physical association) | 0.720 |
| ZNF564 | MDFI | psi-mi:“MI:0915”(physical association) | 0.720 |
| ZNF564 | MID2 | psi-mi:“MI:0915”(physical association) | 0.720 |
| ALAS1 | ZNF564 | psi-mi:“MI:0915”(physical association) | 0.720 |
| ZNF564 | TRIM23 | psi-mi:“MI:0915”(physical association) | 0.720 |
| KRTAP10-7 | ZNF564 | psi-mi:“MI:0915”(physical association) | 0.720 |
| ZNF564 | TRAF1 | psi-mi:“MI:0915”(physical association) | 0.720 |
| MDFI | ZNF564 | psi-mi:“MI:0915”(physical association) | 0.720 |
| MID2 | ZNF564 | psi-mi:“MI:0915”(physical association) | 0.720 |
| ZNF564 | MAGEA1 | psi-mi:“MI:0915”(physical association) | 0.670 |
| MAGEA1 | ZNF564 | psi-mi:“MI:0915”(physical association) | 0.670 |
| ZNF564 | MTUS2 | psi-mi:“MI:0915”(physical association) | 0.670 |
BioGRID (56): ZNF564 (Two-hybrid), ZNF564 (Two-hybrid), ZNF564 (Two-hybrid), ZNF564 (Two-hybrid), ZNF564 (Two-hybrid), ZNF564 (Two-hybrid), ZNF564 (Two-hybrid), ZNF564 (Two-hybrid), ZNF564 (Two-hybrid), ZNF564 (Two-hybrid), ZNF564 (Two-hybrid), ZNF564 (Two-hybrid), ZNF564 (Two-hybrid), ZNF564 (Two-hybrid), ZNF564 (Two-hybrid)
ESM2 similar proteins: A2VDQ7, A6NNF4, A8MQ14, A8MTY0, E9QAG8, O75290, O75373, P08043, P0CJ79, P10751, P17017, P17027, P17039, P35789, P51522, P51523, Q08AN1, Q14585, Q14587, Q3SYV7, Q3ZCX4, Q4R4C7, Q52M93, Q5MCW4, Q5R5U3, Q5R8X1, Q5R9F0, Q5SXM1, Q6ECI4, Q6JLC9, Q6P3V2, Q6P5C7, Q6ZN57, Q86UE3, Q86XN6, Q86YE8, Q8C827, Q8IYB9, Q8N4W9, Q8N7M2
Diamond homologs: A0JPK3, A8MT65, C9JN71, E9QAG8, G3X9G7, O75820, P16373, P16374, P16415, P17017, P17024, P17025, P51815, P52737, Q08AG5, Q08ER8, Q0D2J5, Q15973, Q2M218, Q3KP31, Q3V080, Q494X3, Q4R4C7, Q5MYW4, Q5R9F0, Q5R9S5, Q5REI6, Q5REK1, Q68EA5, Q6P560, Q6P5C7, Q6ZQV5, Q7L945, Q86T29, Q8BGV5, Q8C6P8, Q8IYI8, Q8IZC7, Q8N7K0, Q8N972
SIGNOR signaling
0 interactions.
Enriched among interaction partners
Reactome pathways and GO biological processes over-represented among this gene’s 49 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:
| Pathway | Partners | Fold | FDR |
|---|---|---|---|
| Keratinization | 5 | 12.1× | 6e-03 |
Disease & clinical
Clinical variants and AI predictions
ClinVar
76 variants total. Per-class counts are floors (≥ shown; pagination cap):
| Classification | Count (floor) |
|---|---|
| Pathogenic | 0 |
| Likely pathogenic | 0 |
| Uncertain significance | 74 |
| Likely benign | 1 |
| Benign | 0 |
Top pathogenic / likely-pathogenic (0)
SpliceAI
853 predictions. Top by Δscore:
| Variant | Effect | Δscore |
|---|---|---|
| 19:12527912:GATTT:G | acceptor_gain | 1.0000 |
| 19:12527913:ATTT:A | acceptor_gain | 1.0000 |
| 19:12527914:TTT:T | acceptor_gain | 1.0000 |
| 19:12527915:TT:T | acceptor_gain | 1.0000 |
| 19:12527915:TTC:T | acceptor_loss | 1.0000 |
| 19:12527916:TCTG:T | acceptor_loss | 1.0000 |
| 19:12527917:C:CA | acceptor_loss | 1.0000 |
| 19:12527917:C:CC | acceptor_gain | 1.0000 |
| 19:12527918:T:A | acceptor_loss | 1.0000 |
| 19:12527918:T:C | acceptor_loss | 1.0000 |
| 19:12528564:CCTTA:C | donor_loss | 1.0000 |
| 19:12528565:CTTA:C | donor_loss | 1.0000 |
| 19:12528566:TTA:T | donor_loss | 1.0000 |
| 19:12528567:TA:T | donor_loss | 1.0000 |
| 19:12528568:A:AC | donor_gain | 1.0000 |
| 19:12528568:A:AT | donor_loss | 1.0000 |
| 19:12528569:C:CC | donor_gain | 1.0000 |
| 19:12528569:CCTA:C | donor_gain | 1.0000 |
| 19:12528694:GTCC:G | acceptor_loss | 1.0000 |
| 19:12528695:TCC:T | acceptor_loss | 1.0000 |
| 19:12528697:CTA:C | acceptor_loss | 1.0000 |
| 19:12528697:CTAA:C | acceptor_loss | 1.0000 |
| 19:12528707:CCAAA:C | acceptor_gain | 1.0000 |
| 19:12528708:C:T | acceptor_gain | 1.0000 |
| 19:12528713:A:C | acceptor_gain | 1.0000 |
| 19:12551334:T:TA | donor_gain | 1.0000 |
| 19:12551342:T:TA | donor_gain | 1.0000 |
| 19:12527926:A:AC | acceptor_gain | 0.9900 |
| 19:12527926:A:C | acceptor_gain | 0.9900 |
| 19:12528316:C:A | donor_gain | 0.9900 |
AlphaMissense
3704 scored. Top likely-pathogenic:
| Variant | Protein change | am_pathogenicity |
|---|---|---|
| 19:12526650:G:C | F486L | 0.995 |
| 19:12526650:G:T | F486L | 0.995 |
| 19:12526652:A:G | F486L | 0.995 |
| 19:12526734:A:C | F458L | 0.994 |
| 19:12526734:A:T | F458L | 0.994 |
| 19:12526736:A:G | F458L | 0.994 |
| 19:12526902:A:C | F402L | 0.994 |
| 19:12526902:A:T | F402L | 0.994 |
| 19:12526904:A:G | F402L | 0.994 |
| 19:12527490:A:C | F206L | 0.994 |
| 19:12527490:A:T | F206L | 0.994 |
| 19:12527492:A:G | F206L | 0.994 |
| 19:12526818:G:C | F430L | 0.993 |
| 19:12526818:G:T | F430L | 0.993 |
| 19:12526820:A:G | F430L | 0.993 |
| 19:12527070:G:C | F346L | 0.993 |
| 19:12527070:G:T | F346L | 0.993 |
| 19:12527072:A:G | F346L | 0.993 |
| 19:12527322:A:C | F262L | 0.993 |
| 19:12527322:A:T | F262L | 0.993 |
| 19:12527324:A:G | F262L | 0.993 |
| 19:12527234:A:G | S292P | 0.992 |
| 19:12527238:G:C | F290L | 0.992 |
| 19:12527238:G:T | F290L | 0.992 |
| 19:12527240:A:G | F290L | 0.992 |
| 19:12526611:A:C | H499Q | 0.991 |
| 19:12526611:A:T | H499Q | 0.991 |
| 19:12526695:A:C | H471Q | 0.991 |
| 19:12526695:A:T | H471Q | 0.991 |
| 19:12526863:G:C | H415Q | 0.991 |
dbSNP variants (sampled 300 via entrez): RS1000004564 (19:12550386 T>C,G), RS1000118934 (19:12531291 C>A,T), RS1000161973 (19:12547113 A>G), RS1000407186 (19:12531557 G>A), RS1000621817 (19:12541408 G>A), RS1000678501 (19:12553355 C>A,G,T), RS1000708741 (19:12547397 T>C), RS1001031391 (19:12537548 G>T), RS1001066047 (19:12552179 G>A), RS1001099598 (19:12541451 G>T), RS1001106373 (19:12537332 TTTC>T), RS1001182375 (19:12538496 G>A,T), RS1001187605 (19:12537911 T>G), RS1001318463 (19:12538135 A>G), RS1001344824 (19:12547616 G>C)
Disease associations
OMIM: gene `` | 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.
| Chemical | Actions (top 5) | PubMed papers |
|---|---|---|
| Valproic Acid | affects cotreatment, decreases expression, affects expression | 6 |
| trichostatin A | affects cotreatment, decreases expression | 2 |
| Formaldehyde | decreases expression, increases expression | 2 |
| 3-((6-(2-methoxyphenyl)pyrimidin-4-yl)amino)phenyl)methane sulfonamide | decreases expression | 1 |
| bisphenol F | affects cotreatment, increases expression | 1 |
| TAK-243 | increases sumoylation | 1 |
| methylmercuric chloride | decreases expression | 1 |
| bisphenol A | decreases expression | 1 |
| benzo(e)pyrene | increases methylation | 1 |
| resorcinol | decreases expression | 1 |
| 4-(5-benzo(1,3)dioxol-5-yl-4-pyridin-2-yl-1H-imidazol-2-yl)benzamide | affects cotreatment, decreases expression | 1 |
| dorsomorphin | affects cotreatment, decreases expression | 1 |
| Leflunomide | increases expression | 1 |
| Air Pollutants | decreases expression, increases abundance | 1 |
| Benzo(a)pyrene | decreases methylation | 1 |
| Dexamethasone | affects cotreatment, increases expression | 1 |
| Ethyl Methanesulfonate | increases expression | 1 |
| Indomethacin | affects cotreatment, increases expression | 1 |
| Methapyrilene | increases methylation | 1 |
| Methyl Methanesulfonate | increases expression | 1 |
| Urethane | increases expression | 1 |
| 1-Methyl-3-isobutylxanthine | affects cotreatment, increases expression | 1 |
| Cyclosporine | decreases expression | 1 |
| Copper Sulfate | decreases expression | 1 |
| S-Nitrosoglutathione | decreases expression | 1 |
| Particulate Matter | decreases expression, increases abundance | 1 |
Clinical trials (associated diseases)
0 trials via MONDO — disease-level, not drug-specific.
Related Atlas pages
No linked Atlas pages yet — the cross-entity mesh grows as the corpus expands.