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Rac1 Rac family small GTPase 1 [ Rattus norvegicus (Norway rat) ]

Gene ID: 363875, updated on 11-Apr-2024

Summary

Official Symbol
Rac1provided by RGD
Official Full Name
Rac family small GTPase 1provided by RGD
Primary source
RGD:619755
See related
Ensembl:ENSRNOG00000001068 AllianceGenome:RGD:619755
Gene type
protein coding
RefSeq status
PROVISIONAL
Organism
Rattus norvegicus
Lineage
Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi; Mammalia; Eutheria; Euarchontoglires; Glires; Rodentia; Myomorpha; Muroidea; Muridae; Murinae; Rattus
Summary
Enables several functions, including GTP binding activity; Rho GDP-dissociation inhibitor binding activity; and enzyme binding activity. Involved in several processes, including Rac protein signal transduction; bone resorption; and regulation of cellular biosynthetic process. Located in several cellular components, including Golgi membrane; cytosol; and dendritic spine. Used to study congestive heart failure. Human ortholog(s) of this gene implicated in autosomal dominant mental retardation 48; colon adenocarcinoma; and pancreatic adenocarcinoma. Orthologous to human RAC1 (Rac family small GTPase 1). [provided by Alliance of Genome Resources, Apr 2022]
Expression
Biased expression in Lung (RPKM 434.0), Spleen (RPKM 368.1) and 9 other tissues See more
Orthologs
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Genomic context

Location:
12p11
Exon count:
6
Annotation release Status Assembly Chr Location
RS_2024_02 current GRCr8 (GCF_036323735.1) 12 NC_086030.1 (16150411..16170864)
RS_2023_06 previous assembly mRatBN7.2 (GCF_015227675.2) 12 NC_051347.1 (11037028..11057251)
106 previous assembly Rnor_6.0 (GCF_000001895.5) 12 NC_005111.4 (13090316..13111841)

Chromosome 12 - NC_086030.1Genomic Context describing neighboring genes Neighboring gene ubiquitin specific peptidase 42 Neighboring gene cytohesin 3 Neighboring gene small integral membrane protein 10-like protein 2A Neighboring gene family with sequence similarity 220, member A Neighboring gene diacylglycerol lipase, beta Neighboring gene KDEL endoplasmic reticulum protein retention receptor 2

Genomic regions, transcripts, and products

Expression

  • Project title: A rat RNA-Seq transcriptomic BodyMap across 11 organs and 4 developmental stages
  • Description: 320 RNA samples isolated from 11 organs (adrenal gland, brain, heart, kidney, liver, lung, muscle, spleen, thymus, and testes or uterus) from both sexes of Fischer 344 rats across four developmental stages (2-, 6-, 21-, and 104-weeks-old)
  • BioProject: PRJNA238328
  • Publication: PMID 24510058
  • Analysis date: Mon Jun 6 17:44:12 2016

Bibliography

GeneRIFs: Gene References Into Functions

What's a GeneRIF?

Pathways from PubChem

Interactions

Products Interactant Other Gene Complex Source Pubs Description

General gene information

Markers

Gene Ontology Provided by RGD

Function Evidence Code Pubs
enables ATPase binding IPI
Inferred from Physical Interaction
more info
PubMed 
enables G protein activity ISO
Inferred from Sequence Orthology
more info
 
enables GTP binding IBA
Inferred from Biological aspect of Ancestor
more info
 
enables GTP binding IDA
Inferred from Direct Assay
more info
PubMed 
enables GTP binding IEA
Inferred from Electronic Annotation
more info
 
enables GTP binding ISO
Inferred from Sequence Orthology
more info
 
enables GTP-dependent protein binding ISO
Inferred from Sequence Orthology
more info
 
enables GTPase activity IBA
Inferred from Biological aspect of Ancestor
more info
 
enables GTPase activity IDA
Inferred from Direct Assay
more info
PubMed 
enables GTPase activity IEA
Inferred from Electronic Annotation
more info
 
enables GTPase activity ISO
Inferred from Sequence Orthology
more info
 
enables Rho GDP-dissociation inhibitor binding IPI
Inferred from Physical Interaction
more info
PubMed 
enables enzyme binding ISO
Inferred from Sequence Orthology
more info
 
enables histone deacetylase binding IPI
Inferred from Physical Interaction
more info
PubMed 
enables protein binding IPI
Inferred from Physical Interaction
more info
PubMed 
enables protein kinase binding IBA
Inferred from Biological aspect of Ancestor
more info
 
enables protein kinase binding IEA
Inferred from Electronic Annotation
more info
 
enables protein kinase binding IPI
Inferred from Physical Interaction
more info
PubMed 
enables protein kinase binding ISO
Inferred from Sequence Orthology
more info
 
enables protein-containing complex binding IEA
Inferred from Electronic Annotation
more info
 
enables protein-containing complex binding ISO
Inferred from Sequence Orthology
more info
 
enables small GTPase binding IPI
Inferred from Physical Interaction
more info
PubMed 
enables thioesterase binding IEA
Inferred from Electronic Annotation
more info
 
enables thioesterase binding ISO
Inferred from Sequence Orthology
more info
 
Process Evidence Code Pubs
acts_upstream_of_or_within G protein-coupled receptor signaling pathway ISO
Inferred from Sequence Orthology
more info
 
involved_in Rac protein signal transduction IBA
Inferred from Biological aspect of Ancestor
more info
 
involved_in Rac protein signal transduction IDA
Inferred from Direct Assay
more info
PubMed 
involved_in Rac protein signal transduction ISO
Inferred from Sequence Orthology
more info
 
involved_in Rac protein signal transduction TAS
Traceable Author Statement
more info
PubMed 
acts_upstream_of_or_within Wnt signaling pathway, planar cell polarity pathway ISO
Inferred from Sequence Orthology
more info
 
involved_in actin cytoskeleton organization IDA
Inferred from Direct Assay
more info
PubMed 
involved_in actin cytoskeleton organization IEA
Inferred from Electronic Annotation
more info
 
acts_upstream_of_or_within actin cytoskeleton organization ISO
Inferred from Sequence Orthology
more info
 
involved_in actin filament organization IBA
Inferred from Biological aspect of Ancestor
more info
 
involved_in actin filament organization IMP
Inferred from Mutant Phenotype
more info
PubMed 
acts_upstream_of_or_within actin filament polymerization ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within anatomical structure arrangement ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within angiotensin-activated signaling pathway involved in heart process ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within auditory receptor cell morphogenesis ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within axon guidance ISO
Inferred from Sequence Orthology
more info
 
involved_in bone resorption IDA
Inferred from Direct Assay
more info
PubMed 
acts_upstream_of_or_within cell adhesion ISO
Inferred from Sequence Orthology
more info
 
involved_in cell chemotaxis IBA
Inferred from Biological aspect of Ancestor
more info
 
involved_in cell migration IEA
Inferred from Electronic Annotation
more info
 
acts_upstream_of_or_within cell migration ISO
Inferred from Sequence Orthology
more info
 
involved_in cell migration ISO
Inferred from Sequence Orthology
more info
 
involved_in cell motility ISO
Inferred from Sequence Orthology
more info
 
involved_in cell motility ISS
Inferred from Sequence or Structural Similarity
more info
 
involved_in cell projection assembly IBA
Inferred from Biological aspect of Ancestor
more info
 
acts_upstream_of_or_within cell-cell junction organization ISO
Inferred from Sequence Orthology
more info
 
involved_in cellular response to mechanical stimulus IDA
Inferred from Direct Assay
more info
PubMed 
acts_upstream_of_or_within cerebral cortex GABAergic interneuron development ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within cerebral cortex radially oriented cell migration ISO
Inferred from Sequence Orthology
more info
 
involved_in chemotaxis IMP
Inferred from Mutant Phenotype
more info
PubMed 
acts_upstream_of_or_within cochlea morphogenesis ISO
Inferred from Sequence Orthology
more info
 
involved_in cortical cytoskeleton organization IBA
Inferred from Biological aspect of Ancestor
more info
 
acts_upstream_of_or_within cytoskeleton organization ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within dendrite development ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within dendrite morphogenesis ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within dopaminergic neuron differentiation ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within embryonic olfactory bulb interneuron precursor migration ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within endocytosis ISO
Inferred from Sequence Orthology
more info
 
involved_in engulfment of apoptotic cell IBA
Inferred from Biological aspect of Ancestor
more info
 
involved_in engulfment of apoptotic cell ISO
Inferred from Sequence Orthology
more info
 
involved_in enzyme-linked receptor protein signaling pathway ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within epithelial cell morphogenesis ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within erythrocyte enucleation ISO
Inferred from Sequence Orthology
more info
 
involved_in establishment or maintenance of cell polarity IBA
Inferred from Biological aspect of Ancestor
more info
 
involved_in forebrain development ISO
Inferred from Sequence Orthology
more info
 
involved_in hepatocyte growth factor receptor signaling pathway IEA
Inferred from Electronic Annotation
more info
 
involved_in hepatocyte growth factor receptor signaling pathway ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within homeostasis of number of cells within a tissue ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within hyperosmotic response ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within interneuron migration ISO
Inferred from Sequence Orthology
more info
 
involved_in lamellipodium assembly IEA
Inferred from Electronic Annotation
more info
 
acts_upstream_of_or_within lamellipodium assembly ISO
Inferred from Sequence Orthology
more info
 
involved_in lamellipodium assembly ISO
Inferred from Sequence Orthology
more info
 
involved_in localization ISO
Inferred from Sequence Orthology
more info
 
involved_in localization within membrane IEA
Inferred from Electronic Annotation
more info
 
involved_in localization within membrane ISO
Inferred from Sequence Orthology
more info
 
involved_in mast cell chemotaxis IMP
Inferred from Mutant Phenotype
more info
PubMed 
involved_in midbrain dopaminergic neuron differentiation ISO
Inferred from Sequence Orthology
more info
 
involved_in motor neuron axon guidance IBA
Inferred from Biological aspect of Ancestor
more info
 
involved_in negative regulation of fibroblast migration IEA
Inferred from Electronic Annotation
more info
 
involved_in negative regulation of fibroblast migration ISO
Inferred from Sequence Orthology
more info
 
involved_in negative regulation of interleukin-23 production IEA
Inferred from Electronic Annotation
more info
 
involved_in negative regulation of interleukin-23 production ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within neuron migration ISO
Inferred from Sequence Orthology
more info
 
involved_in neuron migration ISO
Inferred from Sequence Orthology
more info
 
involved_in neuron migration ISS
Inferred from Sequence or Structural Similarity
more info
 
involved_in neuron projection morphogenesis ISO
Inferred from Sequence Orthology
more info
 
involved_in non-canonical Wnt signaling pathway ISO
Inferred from Sequence Orthology
more info
 
involved_in phagocytosis, engulfment ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of DNA replication IMP
Inferred from Mutant Phenotype
more info
PubMed 
acts_upstream_of_or_within positive regulation of actin filament polymerization ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of cell-substrate adhesion ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within positive regulation of dendritic spine development ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of endothelial cell migration IEA
Inferred from Electronic Annotation
more info
 
involved_in positive regulation of endothelial cell migration ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of filopodium assembly IMP
Inferred from Mutant Phenotype
more info
PubMed 
involved_in positive regulation of focal adhesion assembly IEA
Inferred from Electronic Annotation
more info
 
involved_in positive regulation of focal adhesion assembly ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of insulin secretion involved in cellular response to glucose stimulus IMP
Inferred from Mutant Phenotype
more info
PubMed 
involved_in positive regulation of insulin secretion involved in cellular response to glucose stimulus ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of lamellipodium assembly IEA
Inferred from Electronic Annotation
more info
 
involved_in positive regulation of lamellipodium assembly IMP
Inferred from Mutant Phenotype
more info
PubMed 
acts_upstream_of_or_within positive regulation of lamellipodium assembly ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of lamellipodium assembly ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of microtubule polymerization IEA
Inferred from Electronic Annotation
more info
 
involved_in positive regulation of microtubule polymerization ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of neutrophil chemotaxis IEA
Inferred from Electronic Annotation
more info
 
involved_in positive regulation of neutrophil chemotaxis ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of ovarian follicle development ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within positive regulation of phosphatidylinositol 3-kinase/protein kinase B signal transduction ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of protein phosphorylation IEA
Inferred from Electronic Annotation
more info
 
involved_in positive regulation of protein phosphorylation ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of protein phosphorylation ISS
Inferred from Sequence or Structural Similarity
more info
 
involved_in positive regulation of skeletal muscle acetylcholine-gated channel clustering ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of stress fiber assembly IEA
Inferred from Electronic Annotation
more info
 
involved_in positive regulation of stress fiber assembly ISO
Inferred from Sequence Orthology
more info
 
involved_in positive regulation of substrate adhesion-dependent cell spreading IEA
Inferred from Electronic Annotation
more info
 
involved_in positive regulation of substrate adhesion-dependent cell spreading ISO
Inferred from Sequence Orthology
more info
 
involved_in postsynaptic actin cytoskeleton organization EXP
Inferred from Experiment
more info
PubMed 
involved_in postsynaptic actin cytoskeleton organization IDA
Inferred from Direct Assay
more info
PubMed 
involved_in postsynaptic actin cytoskeleton organization IMP
Inferred from Mutant Phenotype
more info
PubMed 
acts_upstream_of_or_within protein localization to plasma membrane ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within regulation of ERK5 cascade ISO
Inferred from Sequence Orthology
more info
 
involved_in regulation of actin cytoskeleton organization IBA
Inferred from Biological aspect of Ancestor
more info
 
involved_in regulation of cell adhesion involved in heart morphogenesis ISO
Inferred from Sequence Orthology
more info
 
involved_in regulation of cell migration ISO
Inferred from Sequence Orthology
more info
 
involved_in regulation of cell migration ISS
Inferred from Sequence or Structural Similarity
more info
 
acts_upstream_of_or_within regulation of cell morphogenesis ISO
Inferred from Sequence Orthology
more info
 
involved_in regulation of cell shape IBA
Inferred from Biological aspect of Ancestor
more info
 
involved_in regulation of cell size IEA
Inferred from Electronic Annotation
more info
 
involved_in regulation of cell size ISO
Inferred from Sequence Orthology
more info
 
involved_in regulation of fibroblast migration ISO
Inferred from Sequence Orthology
more info
 
involved_in regulation of lamellipodium assembly ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within regulation of neuron maturation ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within regulation of neuron migration ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within regulation of neuronal synaptic plasticity ISO
Inferred from Sequence Orthology
more info
 
involved_in regulation of neutrophil migration IBA
Inferred from Biological aspect of Ancestor
more info
 
involved_in regulation of nitric oxide biosynthetic process IGI
Inferred from Genetic Interaction
more info
PubMed 
involved_in regulation of receptor signaling pathway via JAK-STAT ISO
Inferred from Sequence Orthology
more info
 
involved_in regulation of respiratory burst IEA
Inferred from Electronic Annotation
more info
 
involved_in regulation of respiratory burst ISO
Inferred from Sequence Orthology
more info
 
involved_in regulation of stress fiber assembly ISO
Inferred from Sequence Orthology
more info
 
involved_in ruffle assembly IEA
Inferred from Electronic Annotation
more info
 
involved_in ruffle assembly ISO
Inferred from Sequence Orthology
more info
 
acts_upstream_of_or_within ruffle organization ISO
Inferred from Sequence Orthology
more info
 
involved_in semaphorin-plexin signaling pathway IMP
Inferred from Mutant Phenotype
more info
PubMed 
involved_in small GTPase-mediated signal transduction IEA
Inferred from Electronic Annotation
more info
 
acts_upstream_of_or_within small GTPase-mediated signal transduction ISO
Inferred from Sequence Orthology
more info
 
involved_in small GTPase-mediated signal transduction ISO
Inferred from Sequence Orthology
more info
 
involved_in sphingosine-1-phosphate receptor signaling pathway IEA
Inferred from Electronic Annotation
more info
 
involved_in sphingosine-1-phosphate receptor signaling pathway ISO
Inferred from Sequence Orthology
more info
 
involved_in substrate adhesion-dependent cell spreading IEA
Inferred from Electronic Annotation
more info
 
acts_upstream_of_or_within substrate adhesion-dependent cell spreading ISO
Inferred from Sequence Orthology
more info
 
involved_in substrate adhesion-dependent cell spreading ISO
Inferred from Sequence Orthology
more info
 
involved_in substrate adhesion-dependent cell spreading ISS
Inferred from Sequence or Structural Similarity
more info
 
acts_upstream_of_or_within synaptic transmission, GABAergic ISO
Inferred from Sequence Orthology
more info
 
Component Evidence Code Pubs
is_active_in GABA-ergic synapse EXP
Inferred from Experiment
more info
PubMed 
is_active_in GABA-ergic synapse IDA
Inferred from Direct Assay
more info
PubMed 
is_active_in GABA-ergic synapse IMP
Inferred from Mutant Phenotype
more info
PubMed 
located_in Golgi membrane IDA
Inferred from Direct Assay
more info
PubMed 
located_in actin filament IEA
Inferred from Electronic Annotation
more info
 
located_in actin filament ISO
Inferred from Sequence Orthology
more info
 
located_in cell cortex IEA
Inferred from Electronic Annotation
more info
 
located_in cell cortex ISO
Inferred from Sequence Orthology
more info
 
located_in cell projection ISO
Inferred from Sequence Orthology
more info
 
located_in cytoplasm ISO
Inferred from Sequence Orthology
more info
 
located_in cytoplasmic ribonucleoprotein granule IEA
Inferred from Electronic Annotation
more info
 
located_in cytoplasmic ribonucleoprotein granule ISO
Inferred from Sequence Orthology
more info
 
is_active_in cytoplasmic vesicle IBA
Inferred from Biological aspect of Ancestor
more info
 
located_in cytoplasmic vesicle ISO
Inferred from Sequence Orthology
more info
 
is_active_in cytoskeleton IBA
Inferred from Biological aspect of Ancestor
more info
 
located_in cytosol IDA
Inferred from Direct Assay
more info
PubMed 
is_active_in dendritic spine IBA
Inferred from Biological aspect of Ancestor
more info
 
located_in dendritic spine IDA
Inferred from Direct Assay
more info
PubMed 
located_in dendritic spine IEA
Inferred from Electronic Annotation
more info
 
located_in dendritic spine ISO
Inferred from Sequence Orthology
more info
 
located_in early endosome membrane ISO
Inferred from Sequence Orthology
more info
 
located_in glutamatergic synapse IEA
Inferred from Electronic Annotation
more info
 
is_active_in glutamatergic synapse ISO
Inferred from Sequence Orthology
more info
 
located_in kinocilium ISO
Inferred from Sequence Orthology
more info
 
located_in lamellipodium IEA
Inferred from Electronic Annotation
more info
 
located_in lamellipodium ISO
Inferred from Sequence Orthology
more info
 
located_in lamellipodium ISS
Inferred from Sequence or Structural Similarity
more info
 
located_in melanosome IEA
Inferred from Electronic Annotation
more info
 
located_in membrane IDA
Inferred from Direct Assay
more info
PubMed 
located_in membrane ISO
Inferred from Sequence Orthology
more info
 
located_in nucleus IEA
Inferred from Electronic Annotation
more info
 
located_in nucleus ISO
Inferred from Sequence Orthology
more info
 
located_in nucleus ISS
Inferred from Sequence or Structural Similarity
more info
 
located_in pericentriolar material ISO
Inferred from Sequence Orthology
more info
 
located_in phagocytic cup ISO
Inferred from Sequence Orthology
more info
 
is_active_in plasma membrane IBA
Inferred from Biological aspect of Ancestor
more info
 
located_in plasma membrane IEA
Inferred from Electronic Annotation
more info
 
located_in plasma membrane ISO
Inferred from Sequence Orthology
more info
 
located_in plasma membrane TAS
Traceable Author Statement
more info
 
is_active_in postsynapse ISO
Inferred from Sequence Orthology
more info
 
located_in recycling endosome membrane IEA
Inferred from Electronic Annotation
more info
 
located_in recycling endosome membrane ISO
Inferred from Sequence Orthology
more info
 
located_in ruffle membrane IEA
Inferred from Electronic Annotation
more info
 
located_in ruffle membrane ISO
Inferred from Sequence Orthology
more info
 
located_in trans-Golgi network IEA
Inferred from Electronic Annotation
more info
 
located_in trans-Golgi network ISO
Inferred from Sequence Orthology
more info
 

General protein information

Preferred Names
ras-related C3 botulinum toxin substrate 1
Names
p21-Rac1
ras-related C3 botulinum toxin substrate 1 (rho family small GTP binding protein Rac1)
rho family, small GTP binding protein Rac1
NP_599193.1
XP_063127624.1

NCBI Reference Sequences (RefSeq)

NEW Try the new Transcript table

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)

  1. NM_134366.1NP_599193.1  ras-related C3 botulinum toxin substrate 1

    See identical proteins and their annotated locations for NP_599193.1

    Status: PROVISIONAL

    Source sequence(s)
    AY491395
    UniProtKB/Swiss-Prot
    Q6RUV5
    UniProtKB/TrEMBL
    A0A0G2K0X4, A6K1K8
    Related
    ENSRNOP00000075900.2, ENSRNOT00000092558.2
    Conserved Domains (1) summary
    cd01871
    Location:3176
    Rac1_like; Ras-related C3 botulinum toxin substrate 1 (rho family, small GTP binding protein Rac1)-like consists of Rac1, Rac2 and Rac3

RefSeqs of Annotated Genomes: GCF_036323735.1-RS_2024_02

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

Reference GRCr8

Genomic

  1. NC_086030.1 Reference GRCr8

    Range
    16150411..16170864
    Download
    GenBank, FASTA, Sequence Viewer (Graphics)

mRNA and Protein(s)

  1. XM_063271554.1XP_063127624.1  ras-related C3 botulinum toxin substrate 1 isoform X1