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Ras-related C3 botulinum toxin substrate 1



Rattus norvegicus (Rat)
Reviewed-Annotation score: Annotation score: 5 out of 5-Experimental evidence at protein leveli


Plasma membrane-associated small GTPase which cycles between active GTP-bound and inactive GDP-bound states. In its active state, binds to a variety of effector proteins to regulate cellular responses such as secretory processes, phagocytosis of apoptotic cells, epithelial cell polarization and growth-factor induced formation of membrane ruffles. Rac1 p21/rho GDI heterodimer is the active component of the cytosolic factor sigma 1, which is involved in stimulation of the NADPH oxidase activity in macrophages. Essential for the SPATA13-mediated regulation of cell migration and adhesion assembly and disassembly. Stimulates PKN2 kinase activity. In concert with RAB7A, plays a role in regulating the formation of RBs (ruffled borders) in osteoclasts. In glioma cells, promotes cell migration and invasion. In podocytes, promotes nuclear shuttling of NR3C2; this modulation is required for a proper kidney functioning. Required for atypical chemokine receptor ACKR2-induced LIMK1-PAK1-dependent phosphorylation of cofilin (CFL1) and for up-regulation of ACKR2 from endosomal compartment to cell membrane, increasing its efficiency in chemokine uptake and degradation. In synapses, seems to mediate the regulation of F-actin cluster formation performed by SHANK3.5 Publications

Enzyme regulationi

Regulated by guanine nucleotide exchange factors (GEFs) which promote the exchange of bound GDP for free GTP, GTPase activating proteins (GAPs) which increase the GTP hydrolysis activity, and GDP dissociation inhibitors which inhibit the dissociation of the nucleotide from the GTPase. GTP hydrolysis is stimulated by ARHGAP30 (By similarity).By similarity


Feature keyPosition(s)DescriptionActionsGraphical viewLength
Nucleotide bindingi10 – 17GTPBy similarity8
Nucleotide bindingi57 – 61GTPBy similarity5
Nucleotide bindingi115 – 118GTPBy similarity4

GO - Molecular functioni

  • ATPase binding Source: RGD
  • GTPase activity Source: RGD
  • GTP binding Source: RGD
  • histone deacetylase binding Source: RGD
  • protein kinase binding Source: BHF-UCL
  • Rab GTPase binding Source: RGD
  • Rho GDP-dissociation inhibitor binding Source: RGD
  • thioesterase binding Source: Ensembl

GO - Biological processi

  • actin cytoskeleton organization Source: RGD
  • actin filament organization Source: RGD
  • bone resorption Source: RGD
  • cell motility Source: UniProtKB
  • cell proliferation Source: RGD
  • cellular response to mechanical stimulus Source: RGD
  • chemotaxis Source: RGD
  • hepatocyte growth factor receptor signaling pathway Source: Ensembl
  • lamellipodium assembly Source: Ensembl
  • localization within membrane Source: Ensembl
  • mast cell chemotaxis Source: RGD
  • negative regulation of interleukin-23 production Source: Ensembl
  • positive regulation of DNA replication Source: RGD
  • positive regulation of focal adhesion assembly Source: Ensembl
  • positive regulation of lamellipodium assembly Source: RGD
  • positive regulation of microtubule polymerization Source: Ensembl
  • positive regulation of neutrophil chemotaxis Source: Ensembl
  • positive regulation of protein phosphorylation Source: UniProtKB
  • positive regulation of stress fiber assembly Source: Ensembl
  • positive regulation of substrate adhesion-dependent cell spreading Source: Ensembl
  • Rac protein signal transduction Source: SynGO
  • regulation of cell migration Source: UniProtKB
  • regulation of cell size Source: Ensembl
  • regulation of respiratory burst Source: Ensembl
  • ruffle organization Source: Ensembl
  • semaphorin-plexin signaling pathway Source: UniProtKB
  • substrate adhesion-dependent cell spreading Source: Ensembl


LigandGTP-binding, Nucleotide-binding

Enzyme and pathway databases

ReactomeiR-RNO-114604. GPVI-mediated activation cascade.
R-RNO-1433557. Signaling by SCF-KIT.
R-RNO-193648. NRAGE signals death through JNK.
R-RNO-194840. Rho GTPase cycle.
R-RNO-2029482. Regulation of actin dynamics for phagocytic cup formation.
R-RNO-2424491. DAP12 signaling.
R-RNO-2871796. FCERI mediated MAPK activation.
R-RNO-376172. DSCAM interactions.
R-RNO-389359. CD28 dependent Vav1 pathway.
R-RNO-3928662. EPHB-mediated forward signaling.
R-RNO-3928664. Ephrin signaling.
R-RNO-3928665. EPH-ephrin mediated repulsion of cells.
R-RNO-399954. Sema3A PAK dependent Axon repulsion.
R-RNO-4086400. PCP/CE pathway.
R-RNO-416482. G alpha (12/13) signalling events.
R-RNO-416550. Sema4D mediated inhibition of cell attachment and migration.
R-RNO-416572. Sema4D induced cell migration and growth-cone collapse.
R-RNO-418885. DCC mediated attractive signaling.
R-RNO-4420097. VEGFA-VEGFR2 Pathway.
R-RNO-445144. Signal transduction by L1.
R-RNO-5218920. VEGFR2 mediated vascular permeability.
R-RNO-5625970. RHO GTPases activate KTN1.
R-RNO-5626467. RHO GTPases activate IQGAPs.
R-RNO-5627123. RHO GTPases activate PAKs.
R-RNO-5663213. RHO GTPases Activate WASPs and WAVEs.
R-RNO-5668599. RHO GTPases Activate NADPH Oxidases.
R-RNO-5687128. MAPK6/MAPK4 signaling.
R-RNO-6798695. Neutrophil degranulation.
R-RNO-8849471. PTK6 Regulates RHO GTPases, RAS GTPase and MAP kinases.
R-RNO-8875555. MET activates RAP1 and RAC1.
R-RNO-983231. Factors involved in megakaryocyte development and platelet production.

Names & Taxonomyi

Protein namesi
Recommended name:
Ras-related C3 botulinum toxin substrate 1
Alternative name(s):
Gene namesi
OrganismiRattus norvegicus (Rat)
Taxonomic identifieri10116 [NCBI]
Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresGliresRodentiaMyomorphaMuroideaMuridaeMurinaeRattus
  • UP000002494 Componenti: Chromosome 12

Organism-specific databases

RGDi619755. Rac1.

Subcellular locationi

  • Cell membrane; Lipid-anchor; Cytoplasmic side
  • Melanosome By similarity
  • Cytoplasm

  • Note: Inner surface of plasma membrane possibly with attachment requiring prenylation of the C-terminal cysteine (By similarity). Found in the ruffled border (a late endosomal-like compartment in the plasma membrane) of bone-resorbing osteoclasts.By similarity

GO - Cellular componenti

  • actin filament Source: Ensembl
  • cytoplasm Source: RGD
  • cytoplasmic ribonucleoprotein granule Source: Ensembl
  • cytoplasmic vesicle Source: RGD
  • cytosol Source: UniProtKB
  • dendritic spine Source: SynGO
  • extracellular exosome Source: Ensembl
  • extracellular matrix Source: Ensembl
  • focal adhesion Source: Ensembl
  • Golgi membrane Source: RGD
  • lamellipodium Source: Ensembl
  • melanosome Source: UniProtKB-SubCell
  • membrane Source: UniProtKB
  • plasma membrane Source: UniProtKB-SubCell
  • trans-Golgi network Source: Ensembl

Keywords - Cellular componenti

Cell membrane, Cytoplasm, Membrane

Pathology & Biotechi


Feature keyPosition(s)DescriptionActionsGraphical viewLength
Mutagenesisi12G → V: Constitutively active. Increases PAK1 and LIMK1 phosphorylation and NR3C2 nuclear localization in podocytes. 1 Publication1
Mutagenesisi17T → N: Dominant negatif. Reduces NMDA receptor-mediated synaptic currents. 1 Publication1
Mutagenesisi61Q → L: Constitutively active. Interacts with PPP5C. 2 Publications1

PTM / Processingi

Molecule processing

Feature keyPosition(s)DescriptionActionsGraphical viewLength
ChainiPRO_00000420401 – 189Ras-related C3 botulinum toxin substrate 1Add BLAST189
PropeptideiPRO_0000042041190 – 192Removed in mature formBy similarity3

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Cross-linki147Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in ubiquitin)By similarity
Modified residuei189Cysteine methyl esterBy similarity1
Lipidationi189S-geranylgeranyl cysteineBy similarity1

Post-translational modificationi

GTP-bound active form is ubiquitinated by HACE1, leading to its degradation by the proteasome.By similarity

Keywords - PTMi

Isopeptide bond, Lipoprotein, Methylation, Prenylation, Ubl conjugation

Proteomic databases


PTM databases



Tissue specificityi

Osteoclasts.1 Publication

Gene expression databases

ExpressionAtlasiQ6RUV5. baseline and differential.
GenevisibleiQ6RUV5. RN.


Subunit structurei

Interacts with NISCH. Interacts with PIP5K1A. Interacts with the GTP-bound form of RAB7A. Interacts with SRGAP2. Interacts with CYFIP1/SRA-1. Interacts with PLXNB3. Interacts with ARHGDIA; the interaction is induced by SEMA5A, mediated through PLXNB3 and inactivates and stabilizes RAC1. Interacts (GTP-bound form preferentially) with PKN2 (via the REM repeats); the interaction stimulates autophosphorylation and phosphorylation of PKN2. Interacts with the GEF proteins PREX1, RASGRF2, FARP1, FARP2, DOCK1, DOCK2 and DOCK7, which promote the exchange between GDP and GTP, and therefore activate it. Interacts with PARD6A, PARD6B and PARD6G in a GTP-dependent manner. Part of a quaternary complex containing PARD3, some PARD6 protein (PARD6A, PARD6B or PARD6G) and some atypical PKC protein (PRKCI or PRKCZ), which plays a central role in epithelial cell polarization. Found in a trimeric complex composed of DOCK1 and ELMO1, which plays a central role in phagocytosis of apoptotic cells. Interacts with RALBP1 via its effector domain. Interacts with PLXNB1. Probably found in a ternary complex composed of DSCAM, PAK1 and RAC1. Interacts with DSCAM; the interaction requires PAK1. Part of a complex with MAP2K3, MAP3K3, CCM2 and DEF6. Interacts with BAIAP2, BAIAP2L1 and DEF6. Interacts with Y.pseudotuberculosis YPKA and PLCB2. Interacts with NOXA1. Interacts with ARHGEF2. Interacts with TBC1D2. Interacts with UNKL. Interacts with USP6. Interacts with SPATA13. Interacts with ARHGEF16; mediates activation of RAC1 by EPHA2. Interacts with ITGB4. Interacts with S100A8 and calprotectin (S100A8/9). Interacts with PACSIN2. Interacts with ITGB1BP1. Interacts (when active) with PPP5C (via TPR repeats); activates PPP5C phosphatase activity and translocates PPP5C to the cell membrane. Interacts with RAPH1 (via Ras associating and PH domains) (By similarity). Interacts with MTSS1L (via IMD domain); this interaction may be important to potentiate PDGF-induced RAC1 activation (By similarity).By similarity3 Publications

GO - Molecular functioni

  • ATPase binding Source: RGD
  • histone deacetylase binding Source: RGD
  • protein kinase binding Source: BHF-UCL
  • Rab GTPase binding Source: RGD
  • Rho GDP-dissociation inhibitor binding Source: RGD
  • thioesterase binding Source: Ensembl

Protein-protein interaction databases

BioGridi264467. 6 interactors.
IntActiQ6RUV5. 1 interactor.


3D structure databases


Family & Domainsi


Feature keyPosition(s)DescriptionActionsGraphical viewLength
Motifi32 – 40Effector regionSequence analysis9


The effector region mediates interaction with DEF6.By similarity

Sequence similaritiesi

Belongs to the small GTPase superfamily. Rho family.Curated

Phylogenomic databases

eggNOGiKOG0393. Eukaryota.
COG1100. LUCA.

Family and domain databases

InterProiView protein in InterPro
IPR027417. P-loop_NTPase.
IPR005225. Small_GTP-bd_dom.
IPR001806. Small_GTPase.
IPR003578. Small_GTPase_Rho.
PfamiView protein in Pfam
PF00071. Ras. 1 hit.
SUPFAMiSSF52540. SSF52540. 1 hit.
TIGRFAMsiTIGR00231. small_GTP. 1 hit.
PROSITEiView protein in PROSITE
PS51420. RHO. 1 hit.


Sequence statusi: Complete.

Sequence processingi: The displayed sequence is further processed into a mature form.

Q6RUV5-1 [UniParc]FASTAAdd to basket

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        10         20         30         40         50
60 70 80 90 100
110 120 130 140 150
160 170 180 190
Mass (Da):21,450
Last modified:July 5, 2004 - v1

Sequence databases

Select the link destinations:
Links Updated
AY491395 mRNA. Translation: AAR84574.1.
RefSeqiNP_599193.1. NM_134366.1.

Genome annotation databases

EnsembliENSRNOT00000001417; ENSRNOP00000001417; ENSRNOG00000001068.

Similar proteinsi

Links to similar proteins from the UniProt Reference Clusters (UniRef) at 100%, 90% and 50% sequence identity:
100%UniRef100 combines identical sequences and sub-fragments with 11 or more residues from any organism into one UniRef entry.
90%UniRef90 is built by clustering UniRef100 sequences that have at least 90% sequence identity to, and 80% overlap with, the longest sequence (a.k.a seed sequence).
50%UniRef50 is built by clustering UniRef90 seed sequences that have at least 50% sequence identity to, and 80% overlap with, the longest sequence in the cluster.

Entry informationi

Entry nameiRAC1_RAT
AccessioniPrimary (citable) accession number: Q6RUV5
Entry historyiIntegrated into UniProtKB/Swiss-Prot: August 31, 2004
Last sequence update: July 5, 2004
Last modified: July 5, 2017
This is version 134 of the entry and version 1 of the sequence. See complete history.
Entry statusiReviewed (UniProtKB/Swiss-Prot)
Annotation programChordata Protein Annotation Program


Keywords - Technical termi

Complete proteome, Direct protein sequencing, Reference proteome


  1. SIMILARITY comments
    Index of protein domains and families