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Protein

GTP-binding nuclear protein Ran

Gene

Ran

Organism
Rattus norvegicus (Rat)
Status
Reviewed-Annotation score: -Experimental evidence at protein leveli

Functioni

GTPase involved in nucleocytoplasmic transport, participating both to the import and the export from the nucleus of proteins and RNAs. Switches between a cytoplasmic GDP- and a nuclear GTP-bound state by nucleotide exchange and GTP hydrolysis. Nuclear import receptors such as importin beta bind their substrates only in the absence of GTP-bound RAN and release them upon direct interaction with GTP-bound RAN, while export receptors behave in the opposite way. Thereby, RAN controls cargo loading and release by transport receptors in the proper compartment and ensures the directionality of the transport. Interaction with RANBP1 induces a conformation change in the complex formed by XPO1 and RAN that triggers the release of the nuclear export signal of cargo proteins. RAN (GTP-bound form) triggers microtubule assembly at mitotic chromosomes and is required for normal mitotic spindle assembly and chromosome segregation. Required for normal progress through mitosis. The complex with BIRC5/survivin plays a role in mitotic spindle formation by serving as a physical scaffold to help deliver the RAN effector molecule TPX2 to microtubules. Acts as a negative regulator of the kinase activity of VRK1 and VRK2. Enhances AR-mediated transactivation.By similarity

Cofactori

Mg2+By similarityNote: Mg2+ interacts primarily with the phosphate groups of the bound guanine nucleotide.By similarity

Sites

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Binding sitei68GTP; via amide nitrogenBy similarity1
Sitei69Essential for GTP hydrolysisBy similarity1

Regions

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Nucleotide bindingi18 – 25GTPBy similarity8
Nucleotide bindingi36 – 42GTPBy similarity7
Nucleotide bindingi122 – 125GTPBy similarity4
Nucleotide bindingi150 – 152GTPBy similarity3

GO - Molecular functioni

  • dynein intermediate chain binding Source: RGD
  • GDP binding Source: RGD
  • GTPase activity Source: UniProtKB
  • GTP binding Source: RGD
  • importin-alpha family protein binding Source: RGD
  • magnesium ion binding Source: UniProtKB

GO - Biological processi

  • cell division Source: UniProtKB-KW
  • cellular response to mineralocorticoid stimulus Source: RGD
  • GTP metabolic process Source: UniProtKB
  • hippocampus development Source: RGD
  • microtubule cytoskeleton organization Source: RGD
  • mitotic sister chromatid segregation Source: UniProtKB
  • nucleocytoplasmic transport Source: RGD
  • protein export from nucleus Source: RGD
  • protein import into nucleus Source: UniProtKB
  • regulation of protein binding Source: RGD
  • response to organic cyclic compound Source: RGD
  • ribosomal subunit export from nucleus Source: GO_Central
  • snRNA import into nucleus Source: UniProtKB
  • spermatid development Source: RGD

Keywordsi

Biological processCell cycle, Cell division, Mitosis, Protein transport, Transport
LigandGTP-binding, Magnesium, Metal-binding, Nucleotide-binding

Enzyme and pathway databases

ReactomeiR-RNO-1655829 Regulation of cholesterol biosynthesis by SREBP (SREBF)
R-RNO-5578749 Transcriptional regulation by small RNAs

Names & Taxonomyi

Protein namesi
Recommended name:
GTP-binding nuclear protein Ran
Alternative name(s):
GTPase Ran
Ras-like protein TC4
Ras-related nuclear protein
Gene namesi
Name:Ran
OrganismiRattus norvegicus (Rat)
Taxonomic identifieri10116 [NCBI]
Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresGliresRodentiaMyomorphaMuroideaMuridaeMurinaeRattus
Proteomesi
  • UP000002494 Componenti: Chromosome 12

Organism-specific databases

RGDi620367 Ran

Subcellular locationi

Extracellular region or secreted Cytosol Plasma membrane Cytoskeleton Lysosome Endosome Peroxisome ER Golgi apparatus Nucleus Mitochondrion Manual annotation Automatic computational assertionGraphics by Christian Stolte; Source: COMPARTMENTS

Keywords - Cellular componenti

Cytoplasm, Nucleus

PTM / Processingi

Molecule processing

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Initiator methionineiRemovedBy similarity
ChainiPRO_00002086982 – 216GTP-binding nuclear protein RanAdd BLAST215

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Modified residuei2N-acetylalanineBy similarity1
Modified residuei24PhosphothreonineBy similarity1
Modified residuei60N6-acetyllysineBy similarity1
Modified residuei71N6-acetyllysine; alternateBy similarity1
Cross-linki71Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2); alternateBy similarity
Cross-linki71Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in ubiquitin); alternateBy similarity
Modified residuei99N6-acetyllysineBy similarity1
Modified residuei134N6-acetyllysineBy similarity1
Cross-linki152Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2)By similarity
Modified residuei159N6-acetyllysine; alternateBy similarity1
Modified residuei159N6-succinyllysine; alternateBy similarity1

Post-translational modificationi

Acetylation by KAT5 at Lys-134 is increased during mitosis, impairs RANGRF binding and enhances RCC1 binding.By similarity

Keywords - PTMi

Acetylation, Isopeptide bond, Phosphoprotein, Ubl conjugation

Proteomic databases

PaxDbiP62828
PRIDEiP62828

2D gel databases

World-2DPAGEi0004:P62828

PTM databases

CarbonylDBiP62828
iPTMnetiP62828
PhosphoSitePlusiP62828
SwissPalmiP62828

Expressioni

Gene expression databases

BgeeiENSRNOG00000000938
GenevisibleiP62828 RN

Interactioni

Subunit structurei

Monomer. Interacts with RANGAP1, which promotes RAN-mediated GTP hydrolysis. Interacts with KPNB1. Interaction with KPNB1 inhibits RANGAP1-mediated stimulation of GTPase activity. Interacts with RCC1 which promotes the exchange of RAN-bound GDP by GTP. Interaction with KPNB1 inhibits RCC1-mediated exchange of RAN-bound GDP by GTP. Interacts (GTP-bound form) with TNPO1; the interaction is direct. Interacts with KPNB1 and with TNPO1; both inhibit RAN GTPase activity. Interacts (via C-terminus) with RANBP1, which alleviates the inhibition of RAN GTPase activity. Interacts with RANGRF, which promotes the release of bound guanine nucleotide. RANGRF and RCC1 compete for an overlapping binding site on RAN. Identified in a complex with KPNA2 and CSE1L; interaction with RANBP1 mediates dissociation of RAN from this complex. Interaction with both RANBP1 and KPNA2 promotes dissociation of the complex between RAN and KPNB1. Identified in a complex composed of RAN, RANGAP1 and RANBP1. Identified in a complex that contains TNPO1, RAN and RANBP1. Identified in a nuclear export complex with XPO1. Found in a nuclear export complex with RANBP3 and XPO1. Interacts with RANBP2/NUP358. Interaction with RANBP1 or RANBP2 induces a conformation change in the complex formed by XPO1 and RAN that triggers the release of the nuclear export signal of cargo proteins. Component of a nuclear export receptor complex composed of KPNB1, RAN, SNUPN and XPO1 (By similarity). Found in a nuclear export complex with RAN, XPO5 and pre-miRNA (By similarity). Interacts (GTP-bound form) with XPO5 (By similarity). Part of a complex consisting of RANBP9, RAN, DYRK1B and COPS5. Interacts with RANBP9 and RANBP10. Interacts in its GTP-bound form with BIRC5/survivin at S and M phases of the cell cycle. Interacts with TERT; the interaction requires hydrogen peroxide-mediated phosphorylation of TERT and transports TERT to the nucleus. Interacts with MAD2L2. Interacts with VRK1 and VRK3. Interacts with VRK2 (By similarity). Interacts with NEMP1 and KPNB1 (By similarity). Interacts (GDP-bound form) with NUTF2; regulates RAN nuclear import. Interacts with CAPG; mediates CAPG nuclear import. Interacts with NUP153. Interacts with the AR N-terminal poly-Gln region; the interaction with AR is inversely correlated with the poly-Gln length (By similarity).By similarity

GO - Molecular functioni

  • dynein intermediate chain binding Source: RGD
  • importin-alpha family protein binding Source: RGD

Protein-protein interaction databases

BioGridi249998, 3 interactors
IntActiP62828, 1 interactor
STRINGi10116.ENSRNOP00000001247

Structurei

3D structure databases

ProteinModelPortaliP62828
SMRiP62828
ModBaseiSearch...
MobiDBiSearch...

Family & Domainsi

Region

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Regioni211 – 216Interaction with RANBP1By similarity6

Sequence similaritiesi

Belongs to the small GTPase superfamily. Ran family.Curated

Phylogenomic databases

eggNOGiKOG0096 Eukaryota
ENOG410XNRS LUCA
GeneTreeiENSGT00840000129836
HOVERGENiHBG107376
InParanoidiP62828
KOiK07936
OMAiYNVWDTA
OrthoDBiEOG091G0L3X
PhylomeDBiP62828
TreeFamiTF106302

Family and domain databases

InterProiView protein in InterPro
IPR027417 P-loop_NTPase
IPR002041 Ran_GTPase
IPR005225 Small_GTP-bd_dom
IPR001806 Small_GTPase
PANTHERiPTHR24071 PTHR24071, 1 hit
PfamiView protein in Pfam
PF00071 Ras, 1 hit
PRINTSiPR00627 GTPRANTC4
SUPFAMiSSF52540 SSF52540, 1 hit
TIGRFAMsiTIGR00231 small_GTP, 1 hit
PROSITEiView protein in PROSITE
PS51418 RAN, 1 hit

Sequencei

Sequence statusi: Complete.

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

P62828-1 [UniParc]FASTAAdd to basket

« Hide

        10         20         30         40         50
MAAQGEPQVQ FKLVLVGDGG TGKTTFVKRH LTGEFEKKYV ATLGVEVHPL
60 70 80 90 100
VFHTNRGPIK FNVWDTAGQE KFGGLRDGYY IQAQCAIIMF DVTSRVTYKN
110 120 130 140 150
VPNWHRDLVR VCENIPIVLC GNKVDIKDRK VKAKSIVFHR KKNLQYYDIS
160 170 180 190 200
AKSNYNFEKP FLWLARKLIG DPNLEFVAMP ALAPPEVVMD PALAAQYEHD
210
LEVAQTTALP DEDDDL
Length:216
Mass (Da):24,423
Last modified:January 23, 2007 - v3
Checksum:iD5C9B7275C34BCE0
GO

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AF306457 mRNA Translation: AAG33229.1
BC059123 mRNA Translation: AAH59123.1
RefSeqiNP_445891.1, NM_053439.1
XP_006249347.1, XM_006249285.3
UniGeneiRn.107698

Genome annotation databases

EnsembliENSRNOT00000001247; ENSRNOP00000001247; ENSRNOG00000000938
GeneIDi84509
KEGGirno:84509

Similar proteinsi

Entry informationi

Entry nameiRAN_RAT
AccessioniPrimary (citable) accession number: P62828
Secondary accession number(s): P17080
, P28746, P28747, Q9CSP3, Q9CWI7, Q9CZA2, Q9UDJ5, Q9UEU9
Entry historyiIntegrated into UniProtKB/Swiss-Prot: August 16, 2004
Last sequence update: January 23, 2007
Last modified: May 23, 2018
This is version 140 of the entry and version 3 of the sequence. See complete history.
Entry statusiReviewed (UniProtKB/Swiss-Prot)
Annotation programChordata Protein Annotation Program

Miscellaneousi

Keywords - Technical termi

Complete proteome, Direct protein sequencing, Reference proteome

Documents

  1. SIMILARITY comments
    Index of protein domains and families

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