SREBF1 sterol regulatory element binding transcription factor 1 [Homo sapiens (human)] - Gene

Summary

Official Symbol: SREBF1provided by HGNC
Official Full Name: sterol regulatory element binding transcription factor 1provided by HGNC
Primary source: HGNC:HGNC:11289
See related: Ensembl:ENSG00000072310 MIM:184756; AllianceGenome:HGNC:11289
Gene type: protein coding
RefSeq status: REVIEWED
Organism: Homo sapiens
Lineage: Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi; Mammalia; Eutheria; Euarchontoglires; Primates; Haplorrhini; Catarrhini; Hominidae; Homo
Also known as: HMD; IFAP2; SREBP1; bHLHd1
Summary: This gene encodes a basic helix-loop-helix-leucine zipper (bHLH-Zip) transcription factor that binds to the sterol regulatory element-1 (SRE1), which is a motif that is found in the promoter of the low density lipoprotein receptor gene and other genes involved in sterol biosynthesis. The encoded protein is synthesized as a precursor that is initially attached to the nuclear membrane and endoplasmic reticulum. Following cleavage, the mature protein translocates to the nucleus and activates transcription. This cleaveage is inhibited by sterols. This gene is located within the Smith-Magenis syndrome region on chromosome 17. Alternative promoter usage and splicing result in multiple transcript variants, including SREBP-1a and SREBP-1c, which correspond to RefSeq transcript variants 2 and 3, respectively. [provided by RefSeq, Nov 2017]
Expression: Ubiquitous expression in fat (RPKM 35.7), adrenal (RPKM 34.6) and 24 other tissues See more
Orthologs: mouse all
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Genomic context

Location:: 17p11.2

Exon count:: 22

Annotation release	Status	Assembly	Chr	Location
RS_2023_10	current	GRCh38.p14 (GCF_000001405.40)	17	NC_000017.11 (17811334..17836986, complement)
RS_2023_10	current	T2T-CHM13v2.0 (GCF_009914755.1)	17	NC_060941.1 (17757927..17783669, complement)
105.20220307	previous assembly	GRCh37.p13 (GCF_000001405.25)	17	NC_000017.10 (17714648..17740300, complement)

Chromosome 17 - NC_000017.11 Genomic Context describing neighboring genes

Genomic regions, transcripts, and products

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Go to reference sequence details

Genomic Sequence:

Go to nucleotide: Graphics FASTA GenBank

Expression

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See details

Project title: HPA RNA-seq normal tissues
Description: RNA-seq was performed of tissue samples from 95 human individuals representing 27 different tissues in order to determine tissue-specificity of all protein-coding genes
BioProject: PRJEB4337
Publication: PMID 24309898
Analysis date: Wed Apr 4 07:08:55 2018

Bibliography

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GeneRIFs: Gene References Into Functions

What's a GeneRIF?

FAS/SREBP-1c contributes to colorectal carcinogenesis through calcium signaling pathway and cAMP signaling pathway.
Title: FAS/SREBP-1c contributes to colorectal carcinogenesis through calcium signaling pathway and cAMP signaling pathway.
Association of adipocyte size and SREBP-1c in visceral and subcutaneous adipose tissue in non-obese type 2 diabetes mellitus.
Title: Association of adipocyte size and SREBP-1c in visceral and subcutaneous adipose tissue in non-obese type 2 diabetes mellitus.
The Role of SCAP/SREBP as Central Regulators of Lipid Metabolism in Hepatic Steatosis.
Title: The Role of SCAP/SREBP as Central Regulators of Lipid Metabolism in Hepatic Steatosis.
Expression and correlation analysis of silent information regulator 1 (SIRT1), sterol regulatory element-binding protein-1 (SREBP1), and pyroptosis factor in gestational diabetes mellitus.
Title: Expression and correlation analysis of silent information regulator 1 (SIRT1), sterol regulatory element-binding protein-1 (SREBP1), and pyroptosis factor in gestational diabetes mellitus.
EXO1/P53/SREBP1 axis-regulated lipid metabolism promotes prostate cancer progression.
Title: EXO1/P53/SREBP1 axis-regulated lipid metabolism promotes prostate cancer progression.
Exogenous H2 S promotes ubiquitin-mediated degradation of SREBP1 to alleviate diabetic cardiomyopathy via SYVN1 S-sulfhydration.
Title: Exogenous H(2) S promotes ubiquitin-mediated degradation of SREBP1 to alleviate diabetic cardiomyopathy via SYVN1 S-sulfhydration.
Palmitoylation-driven PHF2 ubiquitination remodels lipid metabolism through the SREBP1c axis in hepatocellular carcinoma.
Title: Palmitoylation-driven PHF2 ubiquitination remodels lipid metabolism through the SREBP1c axis in hepatocellular carcinoma.
Recurrent Vascularizing Keratitis in Infants With Hereditary Mucoepithelial Dysplasia Related to SREBF1 Mutation.
Title: Recurrent Vascularizing Keratitis in Infants With Hereditary Mucoepithelial Dysplasia Related to SREBF1 Mutation.
FAM120A couples SREBP-dependent transcription and splicing of lipogenesis enzymes downstream of mTORC1.
Title: FAM120A couples SREBP-dependent transcription and splicing of lipogenesis enzymes downstream of mTORC1.
MIAT shuttled by tumor-secreted exosomes promotes paclitaxel resistance in esophageal cancer cells by activating the TAF1/SREBF1 axis.
Title: MIAT shuttled by tumor-secreted exosomes promotes paclitaxel resistance in esophageal cancer cells by activating the TAF1/SREBF1 axis.

Submit: New GeneRIF Correction See all GeneRIFs (352)

Phenotypes

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Find tests for this gene in the NIH Genetic Testing Registry (GTR)

Review eQTL and phenotype association data in this region using PheGenI

Associated conditions

Description	Tests
Hereditary mucoepithelial dysplasia MedGen: C1274795 OMIM: 158310 GeneReviews: Not available	Compare labs
IFAP syndrome 2 MedGen: C5436607 OMIM: 619016 GeneReviews: Not available	Compare labs

EBI GWAS Catalog

Description
Biological insights from 108 schizophrenia-associated genetic loci. EBI GWAS Catalog EBI GWAS CatalogPubMed
Web-based genome-wide association study identifies two novel loci and a substantial genetic component for Parkinson's disease. EBI GWAS Catalog EBI GWAS CatalogPubMed

Variation

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See variants in ClinVar

See studies and variants in dbVar

See Variation Viewer (GRCh37.p13)

See Variation Viewer (GRCh38)

HIV-1 interactions

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Protein interactions

Protein	Gene	Interaction	Pubs
matrix	gag	HIV-1 MA increases SREBF1 mRNA expression in HepG2 cells and praimary hepatocytes	PubMed

Go to the HIV-1, Human Interaction Database

Interactions

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Products	Interactant	Other Gene	Complex	Source	Pubs	Description

General gene information

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Markers

L29900 (e-PCR)
- UniSTS:50510

GDB:592698 (e-PCR)
- UniSTS:157906

GDB:592707 (e-PCR)
- UniSTS:157907

GDB:592712 (e-PCR)
- UniSTS:157908

GDB:592720 (e-PCR)
- UniSTS:157910

GDB:592722 (e-PCR)
- UniSTS:157911

GDB:592724 (e-PCR)
- UniSTS:157912

GDB:592726 (e-PCR)
- UniSTS:157913

GDB:592728 (e-PCR)
- UniSTS:157914

GDB:592730 (e-PCR)
- UniSTS:157915

GDB:592732 (e-PCR)
- UniSTS:157916

GDB:592734 (e-PCR)
- UniSTS:157917

GDB:592791 (e-PCR)
- UniSTS:157920

GDB:592797 (e-PCR)
- UniSTS:157921

GDB:592800 (e-PCR)
- UniSTS:157922

GDB:592807 (e-PCR)
- UniSTS:157923

GDB:592816 (e-PCR)
- UniSTS:157924

Gene Ontology Provided by GOA

Function	Evidence Code	Pubs
enables DNA binding	ISS Inferred from Sequence or Structural Similarity more info
enables DNA-binding transcription activator activity, RNA polymerase II-specific	IDA Inferred from Direct Assay more info	PubMed
enables DNA-binding transcription factor activity	IDA Inferred from Direct Assay more info	PubMed
enables DNA-binding transcription factor activity, RNA polymerase II-specific	IBA Inferred from Biological aspect of Ancestor more info
enables DNA-binding transcription factor activity, RNA polymerase II-specific	IDA Inferred from Direct Assay more info	PubMed
enables DNA-binding transcription factor activity, RNA polymerase II-specific	IMP Inferred from Mutant Phenotype more info	PubMed
enables DNA-binding transcription factor activity, RNA polymerase II-specific	ISA Inferred from Sequence Alignment more info
enables RNA polymerase II cis-regulatory region sequence-specific DNA binding	IBA Inferred from Biological aspect of Ancestor more info
enables RNA polymerase II transcription regulatory region sequence-specific DNA binding	IDA Inferred from Direct Assay more info	PubMed
enables chromatin binding	IEA Inferred from Electronic Annotation more info
enables nuclear receptor activity	IDA Inferred from Direct Assay more info	PubMed
enables protein binding	IPI Inferred from Physical Interaction more info	PubMed
enables protein dimerization activity	IEA Inferred from Electronic Annotation more info
enables protein kinase binding	IEA Inferred from Electronic Annotation more info
enables sequence-specific double-stranded DNA binding	IDA Inferred from Direct Assay more info	PubMed
enables sterol response element binding	IDA Inferred from Direct Assay more info	PubMed
enables transcription coregulator binding	IPI Inferred from Physical Interaction more info	PubMed

Process	Evidence Code	Pubs
involved_in SREBP signaling pathway	IDA Inferred from Direct Assay more info	PubMed
involved_in SREBP signaling pathway	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in cellular response to fatty acid	IEA Inferred from Electronic Annotation more info
involved_in cellular response to starvation	ISS Inferred from Sequence or Structural Similarity more info	PubMed
involved_in cholesterol biosynthetic process	IEA Inferred from Electronic Annotation more info
involved_in circadian rhythm	IEA Inferred from Electronic Annotation more info
involved_in fat cell differentiation	IEA Inferred from Electronic Annotation more info
involved_in insulin receptor signaling pathway	IEA Inferred from Electronic Annotation more info
involved_in insulin secretion	IEA Inferred from Electronic Annotation more info
involved_in intracellular receptor signaling pathway	IEA Inferred from Electronic Annotation more info
involved_in lipid biosynthetic process	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in lipid biosynthetic process	ISS Inferred from Sequence or Structural Similarity more info
involved_in lipid metabolic process	TAS Traceable Author Statement more info	PubMed
involved_in lung development	IEA Inferred from Electronic Annotation more info
involved_in mRNA transcription by RNA polymerase II	IEA Inferred from Electronic Annotation more info
involved_in negative regulation of insulin secretion	IEA Inferred from Electronic Annotation more info
involved_in negative regulation of transcription by RNA polymerase II	IEA Inferred from Electronic Annotation more info
acts_upstream_of negative regulation of triglyceride metabolic process	IDA Inferred from Direct Assay more info	PubMed
involved_in positive regulation of cholesterol biosynthetic process	IDA Inferred from Direct Assay more info	PubMed
involved_in positive regulation of cholesterol biosynthetic process	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in positive regulation of miRNA transcription	IDA Inferred from Direct Assay more info	PubMed
involved_in positive regulation of transcription by RNA polymerase II	IDA Inferred from Direct Assay more info	PubMed
involved_in positive regulation of transcription by RNA polymerase II	IGI Inferred from Genetic Interaction more info	PubMed
involved_in positive regulation of transcription by RNA polymerase II	TAS Traceable Author Statement more info	PubMed
involved_in positive regulation of triglyceride biosynthetic process	ISS Inferred from Sequence or Structural Similarity more info
involved_in regulation of fatty acid metabolic process	IEA Inferred from Electronic Annotation more info
involved_in regulation of heart rate by chemical signal	IEA Inferred from Electronic Annotation more info
involved_in regulation of lipid storage	IEA Inferred from Electronic Annotation more info
involved_in regulation of mitophagy	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in regulation of protein stability	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in regulation of protein targeting to mitochondrion	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in regulation of transcription by RNA polymerase II	IBA Inferred from Biological aspect of Ancestor more info
involved_in response to cAMP	IEA Inferred from Electronic Annotation more info
involved_in response to ethanol	IEA Inferred from Electronic Annotation more info
involved_in response to food	IEA Inferred from Electronic Annotation more info
involved_in response to fructose	IEA Inferred from Electronic Annotation more info
involved_in response to glucagon	IEA Inferred from Electronic Annotation more info
involved_in response to glucose	IEA Inferred from Electronic Annotation more info
involved_in response to nutrient	IEA Inferred from Electronic Annotation more info
involved_in response to progesterone	IEA Inferred from Electronic Annotation more info
involved_in response to retinoic acid	IEA Inferred from Electronic Annotation more info
involved_in response to xenobiotic stimulus	IEA Inferred from Electronic Annotation more info

Component	Evidence Code	Pubs
located_in ER to Golgi transport vesicle membrane	IEA Inferred from Electronic Annotation more info
located_in Golgi membrane	TAS Traceable Author Statement more info
part_of chromatin	ISA Inferred from Sequence Alignment more info
located_in cytoplasm	ISS Inferred from Sequence or Structural Similarity more info
located_in cytosol	TAS Traceable Author Statement more info
located_in endoplasmic reticulum membrane	TAS Traceable Author Statement more info
located_in nuclear envelope	TAS Traceable Author Statement more info	PubMed
located_in nucleoplasm	TAS Traceable Author Statement 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	IMP Inferred from Mutant Phenotype more info	PubMed

General protein information

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Preferred Names: sterol regulatory element-binding protein 1

Names: class D basic helix-loop-helix protein 1

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.

Genomic

NG_029029.1 RefSeqGene

Range

5026..30678

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GenBank, FASTA, Sequence Viewer (Graphics)

mRNA and Protein(s)

NM_001005291.3 → NP_001005291.1 sterol regulatory element-binding protein 1 isoform 1

See identical proteins and their annotated locations for NP_001005291.1

Status: REVIEWED

Source sequence(s)

AC122129, BC026962, BC057388, BC063281, BQ025047

Consensus CDS

CCDS32583.1

UniProtKB/TrEMBL

B3KR77

Related

ENSP00000348069.4, ENST00000355815.8

Conserved Domains (2) summary

cd00083
Location:351 → 408

HLH; Helix-loop-helix domain, found in specific DNA- binding proteins that act as transcription factors; 60-100 amino acids long. A DNA-binding basic region is followed by two alpha-helices separated by a variable loop region; HLH forms homo- and heterodimers, ...

pfam17040
Location:156 → 356

CBP_CCPA; Cellulose-complementing protein A
NM_001321096.3 → NP_001308025.1 sterol regulatory element-binding protein 1 isoform 3

Status: REVIEWED

Source sequence(s)

AC122129, AK091131, BE208013, DA113045, DB084001

UniProtKB/TrEMBL

B3KR77

Conserved Domains (2) summary

cd00083
Location:297 → 354

HLH; Helix-loop-helix domain, found in specific DNA- binding proteins that act as transcription factors; 60-100 amino acids long. A DNA-binding basic region is followed by two alpha-helices separated by a variable loop region; HLH forms homo- and heterodimers, ...

pfam17040
Location:102 → 302

CBP_CCPA; Cellulose-complementing protein A
NM_001388385.1 → NP_001375314.1 sterol regulatory element-binding protein 1 isoform 4

Status: REVIEWED

Source sequence(s)

AC122129

UniProtKB/TrEMBL

B3KR77

Conserved Domains (3) summary

pfam05109
Location:49 → 320

Herpes_BLLF1; Herpes virus major outer envelope glycoprotein (BLLF1)

cd18921
Location:319 → 393

bHLHzip_SREBP1; basic Helix-Loop-Helix-zipper (bHLHzip) domain found in sterol regulatory element-binding protein 1 (SREBP1) and similar proteins

pfam03154
Location:87 → 210

Atrophin-1; Atrophin-1 family
NM_001388386.1 → NP_001375315.1 sterol regulatory element-binding protein 1 isoform 5

Status: REVIEWED

Source sequence(s)

AC122129

UniProtKB/TrEMBL

B3KR77

Related

ENSP00000379106.2, ENST00000395757.6

Conserved Domains (3) summary

pfam05109
Location:49 → 320

Herpes_BLLF1; Herpes virus major outer envelope glycoprotein (BLLF1)

cd18921
Location:319 → 393

bHLHzip_SREBP1; basic Helix-Loop-Helix-zipper (bHLHzip) domain found in sterol regulatory element-binding protein 1 (SREBP1) and similar proteins

pfam03154
Location:87 → 210

Atrophin-1; Atrophin-1 family
NM_001388387.1 → NP_001375316.1 sterol regulatory element-binding protein 1 isoform 6

Status: REVIEWED

Source sequence(s)

AC122129

UniProtKB/TrEMBL

B3KR77

Conserved Domains (3) summary

pfam05109
Location:49 → 320

Herpes_BLLF1; Herpes virus major outer envelope glycoprotein (BLLF1)

cd18921
Location:319 → 393

bHLHzip_SREBP1; basic Helix-Loop-Helix-zipper (bHLHzip) domain found in sterol regulatory element-binding protein 1 (SREBP1) and similar proteins

pfam03154
Location:87 → 210

Atrophin-1; Atrophin-1 family
NM_001388388.1 → NP_001375317.1 sterol regulatory element-binding protein 1 isoform 7

Status: REVIEWED

Source sequence(s)

AC122129

UniProtKB/TrEMBL

B3KR77

Conserved Domains (3) summary

pfam05109
Location:49 → 320

Herpes_BLLF1; Herpes virus major outer envelope glycoprotein (BLLF1)

pfam03154
Location:87 → 210

Atrophin-1; Atrophin-1 family

cl00081
Location:319 → 378

bHLH_SF; basic Helix Loop Helix (bHLH) domain superfamily
NM_001388389.1 → NP_001375318.1 sterol regulatory element-binding protein 1 isoform 8

Status: REVIEWED

Source sequence(s)

AC122129

UniProtKB/TrEMBL

B3KR77

Conserved Domains (2) summary

cd18921
Location:317 → 391

bHLHzip_SREBP1; basic Helix-Loop-Helix-zipper (bHLHzip) domain found in sterol regulatory element-binding protein 1 (SREBP1) and similar proteins

pfam03154
Location:87 → 210

Atrophin-1; Atrophin-1 family
NM_001388390.1 → NP_001375319.1 sterol regulatory element-binding protein 1 isoform 9

Status: REVIEWED

Source sequence(s)

AC122129

UniProtKB/TrEMBL

B3KR77

Conserved Domains (3) summary

pfam05109
Location:49 → 320

Herpes_BLLF1; Herpes virus major outer envelope glycoprotein (BLLF1)

cd18921
Location:319 → 387

bHLHzip_SREBP1; basic Helix-Loop-Helix-zipper (bHLHzip) domain found in sterol regulatory element-binding protein 1 (SREBP1) and similar proteins

pfam03154
Location:87 → 210

Atrophin-1; Atrophin-1 family
NM_001388391.1 → NP_001375320.1 sterol regulatory element-binding protein 1 isoform 10

Status: REVIEWED

Source sequence(s)

AC122129

UniProtKB/TrEMBL

B3KR77

Conserved Domains (3) summary

pfam05109
Location:49 → 320

Herpes_BLLF1; Herpes virus major outer envelope glycoprotein (BLLF1)

cd18921
Location:319 → 384

bHLHzip_SREBP1; basic Helix-Loop-Helix-zipper (bHLHzip) domain found in sterol regulatory element-binding protein 1 (SREBP1) and similar proteins

pfam03154
Location:87 → 210

Atrophin-1; Atrophin-1 family
NM_001388392.1 → NP_001375321.1 sterol regulatory element-binding protein 1 isoform 11

Status: REVIEWED

Source sequence(s)

AC122129

UniProtKB/TrEMBL

B3KR77

Conserved Domains (2) summary

pfam03154
Location:87 → 210

Atrophin-1; Atrophin-1 family

cl00081
Location:267 → 319

bHLH_SF; basic Helix Loop Helix (bHLH) domain superfamily
NM_001388393.1 → NP_001375322.1 sterol regulatory element-binding protein 1 isoform 12

Status: REVIEWED

Source sequence(s)

AC122129

UniProtKB/TrEMBL

B3KR77

Conserved Domains (3) summary

pfam05109
Location:49 → 320

Herpes_BLLF1; Herpes virus major outer envelope glycoprotein (BLLF1)

cd18921
Location:319 → 393

bHLHzip_SREBP1; basic Helix-Loop-Helix-zipper (bHLHzip) domain found in sterol regulatory element-binding protein 1 (SREBP1) and similar proteins

pfam03154
Location:87 → 210

Atrophin-1; Atrophin-1 family
NM_001388394.1 → NP_001375323.1 sterol regulatory element-binding protein 1 isoform 13

Status: REVIEWED

Source sequence(s)

AC122129

UniProtKB/TrEMBL

B3KXZ7

Conserved Domains (1) summary

cd18921
Location:175 → 249

bHLHzip_SREBP1; basic Helix-Loop-Helix-zipper (bHLHzip) domain found in sterol regulatory element-binding protein 1 (SREBP1) and similar proteins
NM_004176.5 → NP_004167.3 sterol regulatory element-binding protein 1 isoform 2

See identical proteins and their annotated locations for NP_004167.3

Status: REVIEWED

Source sequence(s)

AC122129, BC026962, BC057388, BC063281, BQ025047

Consensus CDS

CCDS11189.1

UniProtKB/Swiss-Prot

B0I4X3, B0I4X4, D3DXC4, P36956, Q16062, Q59F52, Q6P4R7, Q6PFW7, Q6PJ36, Q8TAK9

UniProtKB/TrEMBL

B3KR77

Related

ENSP00000261646.5, ENST00000261646.11

Conserved Domains (3) summary

cd00083
Location:321 → 378

HLH; Helix-loop-helix domain, found in specific DNA- binding proteins that act as transcription factors; 60-100 amino acids long. A DNA-binding basic region is followed by two alpha-helices separated by a variable loop region; HLH forms homo- and heterodimers, ...

cl25496
Location:49 → 320

Herpes_BLLF1; Herpes virus major outer envelope glycoprotein (BLLF1)

cl27975
Location:87 → 212

EBV-NA3; Epstein-Barr virus nuclear antigen 3 (EBNA-3)

RNA

NR_170943.1 RNA Sequence

Status: REVIEWED

Source sequence(s)

AC122129
NR_170944.1 RNA Sequence

Status: REVIEWED

Source sequence(s)

AC122129
NR_170945.1 RNA Sequence

Status: REVIEWED

Source sequence(s)

AC122129
NR_170990.1 RNA Sequence

Status: REVIEWED

Source sequence(s)

AC122129

RefSeqs of Annotated Genomes: GCF_000001405.40-RS_2023_10

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

Reference GRCh38.p14 Primary Assembly

Genomic

NC_000017.11 Reference GRCh38.p14 Primary Assembly

Range

17811334..17836986 complement

Download

GenBank, FASTA, Sequence Viewer (Graphics)

mRNA and Protein(s)

XM_047436580.1 → XP_047292536.1 sterol regulatory element-binding protein 1 isoform X1
XM_047436581.1 → XP_047292537.1 sterol regulatory element-binding protein 1 isoform X2

Alternate T2T-CHM13v2.0

Genomic

NC_060941.1 Alternate T2T-CHM13v2.0

Range

17757927..17783669 complement

Download

GenBank, FASTA, Sequence Viewer (Graphics)

mRNA and Protein(s)

XM_054316989.1 → XP_054172964.1 sterol regulatory element-binding protein 1 isoform X1
XM_054316990.1 → XP_054172965.1 sterol regulatory element-binding protein 1 isoform X2