Chd5 chromodomain helicase DNA binding protein 5 [Mus musculus (house mouse)] - Gene

Summary

Official Symbol: Chd5provided by MGI
Official Full Name: chromodomain helicase DNA binding protein 5provided by MGI
Primary source: MGI:MGI:3036258
See related: Ensembl:ENSMUSG00000005045 AllianceGenome:MGI:3036258
Gene type: protein coding
RefSeq status: VALIDATED
Organism: Mus musculus
Lineage: Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi; Mammalia; Eutheria; Euarchontoglires; Glires; Rodentia; Myomorpha; Muroidea; Muridae; Murinae; Mus; Mus
Also known as: CHD-5; 4930532L22Rik; B230399N07Rik
Summary: Enables H3K27me3 modified histone binding activity and histone deacetylase binding activity. Involved in several processes, including negative regulation of transcription by RNA polymerase II; positive regulation of signal transduction by p53 class mediator; and sperm DNA condensation. Located in heterochromatin and nucleus. Part of NuRD complex. Is expressed in central nervous system and dorsal root ganglion. Orthologous to human CHD5 (chromodomain helicase DNA binding protein 5). [provided by Alliance of Genome Resources, Nov 2024]
Expression: Biased expression in testis adult (RPKM 38.0), frontal lobe adult (RPKM 16.9) and 5 other tissues See more
Orthologs: human all
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Genomic context

Location:: 4 E2; 4 83.04 cM

Exon count:: 43

Annotation release	Status	Assembly	Chr	Location
RS_2024_02	current	GRCm39 (GCF_000001635.27)	4	NC_000070.7 (152423103..152474651)
108.20200622	previous assembly	GRCm38.p6 (GCF_000001635.26)	4	NC_000070.6 (152338646..152390194)

Chromosome 4 - NC_000070.7 Genomic Context describing neighboring genes

Genomic regions, transcripts, and products

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Go to reference sequence details

Genomic Sequence:

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Expression

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See details

Project title: Mouse ENCODE transcriptome data Mouse ENCODE transcriptome data
Description: RNA profiling data sets generated by the Mouse ENCODE project.
BioProject: PRJNA66167
Publication: PMID 25409824
Analysis date: n/a

Bibliography

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GeneRIFs: Gene References Into Functions

What's a GeneRIF?

Chd5 Regulates the Transcription Factor Six3 to Promote Neuronal Differentiation.
Title: Chd5 Regulates the Transcription Factor Six3 to Promote Neuronal Differentiation.
Pyramidal neurons cultured from Chd5(-/-) cortex displayed alterations in dendritic morphology. Paralleling autism spectrum disorder nosology, Chd5(-/-) mice exhibited abnormal sociocommunicative behavior and a strong preference for familiarity. Chd5(-/-) mice further showed deficits in responding to the distress of a conspecific, a mouse homolog of empathy.
Title: Mice lacking the chromodomain helicase DNA-binding 5 chromatin remodeler display autism-like characteristics.
Chd5 represses murine endogenous retrovirus-L (MuERV-L/MERVL), an endogenous retrovirus-derived retrotransposon, by regulating H3K27me3 and H3.1/H3.2 function.
Title: Chd5 Regulates MuERV-L/MERVL Expression in Mouse Embryonic Stem Cells Via H3K27me3 Modification and Histone H3.1/H3.2.
Chd5 is a multi-faceted mediator of histone-to-protamine replacement and is important in the cascade of molecular events underlying this process of extensive chromatin remodelling.
Title: Chd5 orchestrates chromatin remodelling during sperm development.
Our data indicate that CHD5 is required for normal spermiogenesis, especially for spermatid chromatin condensation.
Title: CHD5 is required for spermiogenesis and chromatin condensation.
Chd5 expression is retained at high levels in differentiated neurons of the adult mouse.
Title: The tumor suppressor Chd5 is induced during neuronal differentiation in the developing mouse brain.
These findings provide insights into the regulatory role of CHD5 during neurogenesis and suggest how inactivation of this candidate tumor suppressor might contribute to neuroblastoma.
Title: CHD5 is required for neurogenesis and has a dual role in facilitating gene expression and polycomb gene repression.
Chd5 binds the unmodified N terminus of H3 through its tandem plant homeodomains (PHDs).
Title: Chd5 requires PHD-mediated histone 3 binding for tumor suppression.
CHD5 regulates expression of neuronal genes, cell cycle genes and remodeling genes, and is linked to regulation of genes implicated in aging and Alzheimer's disease.
Title: CHD5, a brain-specific paralog of Mi2 chromatin remodeling enzymes, regulates expression of neuronal genes.
the association of between miR-211-regulated CHD5 expression and CHD5 function in colorectal tumorigenesis
Title: MicroRNA-211 expression promotes colorectal cancer cell growth in vitro and in vivo by targeting tumor suppressor CHD5.

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Variation

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Alleles

Alleles of this type are documented at Mouse Genome Informatics (MGI)

Endonuclease-mediated (2)
Targeted (5) 1 citation

Interactions

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Products	Interactant	Other Gene	Complex	Source	Pubs	Description

General gene information

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Markers

RH132291 (e-PCR)
- UniSTS:215574

AW060752 (e-PCR)
- UniSTS:179119

Gene Ontology Provided by MGI

Function	Evidence Code	Pubs
enables ATP binding	IEA Inferred from Electronic Annotation more info
enables ATP hydrolysis activity	IBA Inferred from Biological aspect of Ancestor more info
enables ATP hydrolysis activity	IEA Inferred from Electronic Annotation more info
enables ATP-dependent chromatin remodeler activity	IBA Inferred from Biological aspect of Ancestor more info
enables ATP-dependent chromatin remodeler activity	IEA Inferred from Electronic Annotation more info
enables DNA binding	IBA Inferred from Biological aspect of Ancestor more info
enables DNA binding	IEA Inferred from Electronic Annotation more info
enables H3K27me3 modified histone binding	IDA Inferred from Direct Assay more info	PubMed
enables H3K27me3 modified histone binding	ISO Inferred from Sequence Orthology more info
enables chromatin binding	IBA Inferred from Biological aspect of Ancestor more info
enables chromatin binding	ISO Inferred from Sequence Orthology more info
enables helicase activity	IEA Inferred from Electronic Annotation more info
enables histone binding	IBA Inferred from Biological aspect of Ancestor more info
enables histone binding	IDA Inferred from Direct Assay more info	PubMed
enables histone deacetylase binding	IPI Inferred from Physical Interaction more info	PubMed
enables metal ion binding	IEA Inferred from Electronic Annotation more info
enables protein binding	IPI Inferred from Physical Interaction more info	PubMed

Process	Evidence Code	Pubs
involved_in cerebral cortex neuron differentiation	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in chromatin remodeling	IBA Inferred from Biological aspect of Ancestor more info
involved_in chromatin remodeling	ISO Inferred from Sequence Orthology more info
involved_in chromatin remodeling	ISS Inferred from Sequence or Structural Similarity more info
involved_in negative regulation of cell population proliferation	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in negative regulation of transcription by RNA polymerase II	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in positive regulation of signal transduction by p53 class mediator	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in regulation of cell differentiation	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in sperm DNA condensation	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in transcription by RNA polymerase II	ISO Inferred from Sequence Orthology more info

Component	Evidence Code	Pubs
part_of NuRD complex	IBA Inferred from Biological aspect of Ancestor more info
part_of NuRD complex	IDA Inferred from Direct Assay more info	PubMed
located_in chromatin	IBA Inferred from Biological aspect of Ancestor more info
located_in chromatin	ISO Inferred from Sequence Orthology more info
located_in cytosol	ISO Inferred from Sequence Orthology more info
located_in heterochromatin	IDA Inferred from Direct Assay more info	PubMed
located_in nuclear speck	ISO Inferred from Sequence Orthology more info
located_in nucleoplasm	ISO Inferred from Sequence Orthology more info
is_active_in nucleus	IBA Inferred from Biological aspect of Ancestor more info
located_in nucleus	IDA Inferred from Direct Assay more info	PubMed
located_in nucleus	ISO Inferred from Sequence Orthology more info

General protein information

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Preferred Names: chromodomain-helicase-DNA-binding protein 5

Names: ATP-dependent helicase CHD5

NP_001074845.1

EC 3.6.4.12

NP_001356172.1

EC 3.6.4.12

NP_083492.2

EC 3.6.4.12

XP_006538992.1

EC 3.6.4.12

XP_006538993.1

EC 3.6.4.12

XP_006538994.1

EC 3.6.4.12

XP_006538996.1

EC 3.6.4.12

XP_006538998.1

EC 3.6.4.12

XP_017175733.1

EC 3.6.4.12

XP_030109449.1

EC 3.6.4.12

NCBI Reference Sequences (RefSeq)

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RefSeqs maintained independently of Annotated Genomes

These reference sequences exist independently of genome builds. Explain

These reference sequences are curated independently of the genome annotation cycle, so their versions may not match the RefSeq versions in the current genome build. Identify version mismatches by comparing the version of the RefSeq in this section to the one reported in Genomic regions, transcripts, and products above.

mRNA and Protein(s)

NM_001081376.1 → NP_001074845.1 chromodomain-helicase-DNA-binding protein 5 isoform 1

Status: VALIDATED

Description

Transcript Variant: This variant (1) represents the longer transcript and encodes the longer isoform (1).

Source sequence(s)

AL611985

Consensus CDS

CCDS38984.1

UniProtKB/TrEMBL

E9PYL1

Related

ENSMUSP00000030775.6, ENSMUST00000030775.12

Conserved Domains (10) summary

smart00298
Location:594 → 648

CHROMO; Chromatin organization modifier domain

cd00046
Location:722 → 880

DEXDc; DEAD-like helicases superfamily. A diverse family of proteins involved in ATP-dependent RNA or DNA unwinding. This domain contains the ATP-binding region.

cd15531
Location:347 → 389

PHD1_CHD_II; PHD finger 1 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd15532
Location:420 → 462

PHD2_CHD_II; PHD finger 2 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

pfam00176
Location:705 → 1001

SNF2_N; SNF2 family N-terminal domain

pfam00271
Location:1025 → 1140

Helicase_C; Helicase conserved C-terminal domain

pfam06461
Location:1390 → 1531

DUF1086; Domain of Unknown Function (DUF1086)

pfam06465
Location:1300 → 1360

DUF1087; Domain of Unknown Function (DUF1087)

pfam08073
Location:153 → 202

CHDNT; CHDNT (NUC034) domain

pfam08074
Location:1730 → 1901

CHDCT2; CHDCT2 (NUC038) domain
NM_001369243.1 → NP_001356172.1 chromodomain-helicase-DNA-binding protein 5 isoform 3

Status: VALIDATED

Source sequence(s)

AL611985

Consensus CDS

CCDS89866.1

UniProtKB/Swiss-Prot

A2A8L1, Q6ZQB2

Related

ENSMUSP00000005175.5, ENSMUST00000005175.5

Conserved Domains (10) summary

PLN03142
Location:700 → 1218

PLN03142; Probable chromatin-remodeling complex ATPase chain; Provisional

PRK13108
Location:1532 → 1684

PRK13108; prolipoprotein diacylglyceryl transferase; Reviewed

cd15531
Location:347 → 389

PHD1_CHD_II; PHD finger 1 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd15532
Location:420 → 462

PHD2_CHD_II; PHD finger 2 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

pfam06461
Location:1390 → 1531

DUF1086; Domain of Unknown Function (DUF1086)

pfam08074
Location:1730 → 1901

CHDCT2; CHDCT2 (NUC038) domain

pfam06465
Location:1300 → 1359

DUF1087; Domain of Unknown Function (DUF1087)

pfam08073
Location:153 → 203

CHDNT; CHDNT (NUC034) domain

cd18662
Location:591 → 644

CD2_tandem_CHD3-4_like; repeat 2 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

cd18667
Location:467 → 546

CD1_tandem_CHD3-4_like; repeat 1 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins
NM_029216.2 → NP_083492.2 chromodomain-helicase-DNA-binding protein 5 isoform 2

Status: VALIDATED

Description

Transcript Variant: This variant (2) lacks an in-frame exon, compared to variant 1. It encodes isoform 2 which is shorter than isoform 1.

Source sequence(s)

AL611985

Consensus CDS

CCDS51394.1

UniProtKB/TrEMBL

E9PYU4

Related

ENSMUSP00000132600.2, ENSMUST00000164662.8

Conserved Domains (10) summary

smart00298
Location:594 → 648

CHROMO; Chromatin organization modifier domain

cd00046
Location:722 → 880

DEXDc; DEAD-like helicases superfamily. A diverse family of proteins involved in ATP-dependent RNA or DNA unwinding. This domain contains the ATP-binding region.

cd15531
Location:347 → 389

PHD1_CHD_II; PHD finger 1 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd15532
Location:420 → 462

PHD2_CHD_II; PHD finger 2 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

pfam00176
Location:705 → 1001

SNF2_N; SNF2 family N-terminal domain

pfam00271
Location:1025 → 1140

Helicase_C; Helicase conserved C-terminal domain

pfam06461
Location:1353 → 1494

DUF1086; Domain of Unknown Function (DUF1086)

pfam06465
Location:1263 → 1323

DUF1087; Domain of Unknown Function (DUF1087)

pfam08073
Location:153 → 202

CHDNT; CHDNT (NUC034) domain

pfam08074
Location:1693 → 1864

CHDCT2; CHDCT2 (NUC038) domain

RefSeqs of Annotated Genomes: GCF_000001635.27-RS_2024_02

The following sections contain reference sequences that belong to a specific genome build. Explain

Reference GRCm39 C57BL/6J

Genomic

NC_000070.7 Reference GRCm39 C57BL/6J

Range

152423103..152474651

Download

GenBank, FASTA, Sequence Viewer (Graphics)

mRNA and Protein(s)

XM_006538929.2 → XP_006538992.1 chromodomain-helicase-DNA-binding protein 5 isoform X1

Conserved Domains (10) summary

PLN03142
Location:700 → 1218

PLN03142; Probable chromatin-remodeling complex ATPase chain; Provisional

PRK13108
Location:1535 → 1687

PRK13108; prolipoprotein diacylglyceryl transferase; Reviewed

cd15531
Location:347 → 389

PHD1_CHD_II; PHD finger 1 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd15532
Location:420 → 462

PHD2_CHD_II; PHD finger 2 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd18662
Location:591 → 644

CD2_tandem_CHD3-4_like; repeat 2 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

cd18667
Location:467 → 546

CD1_tandem_CHD3-4_like; repeat 1 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

pfam06461
Location:1393 → 1534

DUF1086; Domain of Unknown Function (DUF1086)

pfam06465
Location:1300 → 1359

DUF1087; Domain of Unknown Function (DUF1087)

pfam08073
Location:148 → 203

CHDNT; CHDNT (NUC034) domain

pfam08074
Location:1733 → 1904

CHDCT2; CHDCT2 (NUC038) domain
XM_006538933.2 → XP_006538996.1 chromodomain-helicase-DNA-binding protein 5 isoform X5

Conserved Domains (10) summary

PLN03142
Location:700 → 1240

PLN03142; Probable chromatin-remodeling complex ATPase chain; Provisional

PRK13108
Location:1498 → 1650

PRK13108; prolipoprotein diacylglyceryl transferase; Reviewed

cd15531
Location:347 → 389

PHD1_CHD_II; PHD finger 1 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd15532
Location:420 → 462

PHD2_CHD_II; PHD finger 2 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd18662
Location:591 → 644

CD2_tandem_CHD3-4_like; repeat 2 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

cd18667
Location:467 → 546

CD1_tandem_CHD3-4_like; repeat 1 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

pfam06461
Location:1356 → 1497

DUF1086; Domain of Unknown Function (DUF1086)

pfam06465
Location:1263 → 1322

DUF1087; Domain of Unknown Function (DUF1087)

pfam08073
Location:148 → 203

CHDNT; CHDNT (NUC034) domain

pfam08074
Location:1696 → 1867

CHDCT2; CHDCT2 (NUC038) domain
XM_006538930.2 → XP_006538993.1 chromodomain-helicase-DNA-binding protein 5 isoform X2

Conserved Domains (10) summary

PLN03142
Location:700 → 1217

PLN03142; Probable chromatin-remodeling complex ATPase chain; Provisional

PRK13108
Location:1534 → 1686

PRK13108; prolipoprotein diacylglyceryl transferase; Reviewed

cd15531
Location:347 → 389

PHD1_CHD_II; PHD finger 1 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd15532
Location:420 → 462

PHD2_CHD_II; PHD finger 2 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd18662
Location:591 → 644

CD2_tandem_CHD3-4_like; repeat 2 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

cd18667
Location:467 → 546

CD1_tandem_CHD3-4_like; repeat 1 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

pfam06461
Location:1392 → 1533

DUF1086; Domain of Unknown Function (DUF1086)

pfam06465
Location:1299 → 1358

DUF1087; Domain of Unknown Function (DUF1087)

pfam08073
Location:148 → 203

CHDNT; CHDNT (NUC034) domain

pfam08074
Location:1732 → 1903

CHDCT2; CHDCT2 (NUC038) domain
XM_017320244.2 → XP_017175733.1 chromodomain-helicase-DNA-binding protein 5 isoform X3

Conserved Domains (10) summary

PLN03142
Location:700 → 1217

PLN03142; Probable chromatin-remodeling complex ATPase chain; Provisional

PRK13108
Location:1531 → 1683

PRK13108; prolipoprotein diacylglyceryl transferase; Reviewed

cd15531
Location:347 → 389

PHD1_CHD_II; PHD finger 1 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd15532
Location:420 → 462

PHD2_CHD_II; PHD finger 2 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd18662
Location:591 → 644

CD2_tandem_CHD3-4_like; repeat 2 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

cd18667
Location:467 → 546

CD1_tandem_CHD3-4_like; repeat 1 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

pfam06461
Location:1389 → 1530

DUF1086; Domain of Unknown Function (DUF1086)

pfam06465
Location:1299 → 1358

DUF1087; Domain of Unknown Function (DUF1087)

pfam08073
Location:148 → 203

CHDNT; CHDNT (NUC034) domain

pfam08074
Location:1729 → 1900

CHDCT2; CHDCT2 (NUC038) domain
XM_006538931.2 → XP_006538994.1 chromodomain-helicase-DNA-binding protein 5 isoform X4

Conserved Domains (10) summary

PLN03142
Location:700 → 1218

PLN03142; Probable chromatin-remodeling complex ATPase chain; Provisional

PRK13108
Location:1535 → 1687

PRK13108; prolipoprotein diacylglyceryl transferase; Reviewed

cd15531
Location:347 → 389

PHD1_CHD_II; PHD finger 1 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd15532
Location:420 → 462

PHD2_CHD_II; PHD finger 2 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd18662
Location:591 → 644

CD2_tandem_CHD3-4_like; repeat 2 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

cd18667
Location:467 → 546

CD1_tandem_CHD3-4_like; repeat 1 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

pfam06461
Location:1393 → 1534

DUF1086; Domain of Unknown Function (DUF1086)

pfam06465
Location:1300 → 1359

DUF1087; Domain of Unknown Function (DUF1087)

pfam08073
Location:148 → 203

CHDNT; CHDNT (NUC034) domain

pfam08074
Location:1733 → 1904

CHDCT2; CHDCT2 (NUC038) domain
XM_006538935.2 → XP_006538998.1 chromodomain-helicase-DNA-binding protein 5 isoform X7

Conserved Domains (10) summary

PLN03142
Location:700 → 1218

PLN03142; Probable chromatin-remodeling complex ATPase chain; Provisional

PRK13108
Location:1535 → 1687

PRK13108; prolipoprotein diacylglyceryl transferase; Reviewed

cd15531
Location:347 → 389

PHD1_CHD_II; PHD finger 1 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd15532
Location:420 → 462

PHD2_CHD_II; PHD finger 2 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd18662
Location:591 → 644

CD2_tandem_CHD3-4_like; repeat 2 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

cd18667
Location:467 → 546

CD1_tandem_CHD3-4_like; repeat 1 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

pfam06461
Location:1393 → 1534

DUF1086; Domain of Unknown Function (DUF1086)

pfam06465
Location:1300 → 1359

DUF1087; Domain of Unknown Function (DUF1087)

pfam08073
Location:148 → 203

CHDNT; CHDNT (NUC034) domain

pfam08074
Location:1733 → 1860

CHDCT2; CHDCT2 (NUC038) domain
XM_030253589.1 → XP_030109449.1 chromodomain-helicase-DNA-binding protein 5 isoform X6

Conserved Domains (10) summary

PLN03142
Location:626 → 1144

PLN03142; Probable chromatin-remodeling complex ATPase chain; Provisional

PRK13108
Location:1461 → 1613

PRK13108; prolipoprotein diacylglyceryl transferase; Reviewed

cd15531
Location:273 → 315

PHD1_CHD_II; PHD finger 1 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd15532
Location:346 → 388

PHD2_CHD_II; PHD finger 2 found in class II Chromodomain-Helicase-DNA binding (CHD) proteins

cd18662
Location:517 → 570

CD2_tandem_CHD3-4_like; repeat 2 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

cd18667
Location:393 → 472

CD1_tandem_CHD3-4_like; repeat 1 of the paired tandem chromodomains of chromodomain helicase DNA-binding protein 3 and 4, and similar proteins

pfam06461
Location:1319 → 1460

DUF1086; Domain of Unknown Function (DUF1086)

pfam06465
Location:1226 → 1285

DUF1087; Domain of Unknown Function (DUF1087)

pfam08073
Location:74 → 129

CHDNT; CHDNT (NUC034) domain

pfam08074
Location:1659 → 1830

CHDCT2; CHDCT2 (NUC038) domain