P55060 (XPO2_HUMAN) Reviewed, UniProtKB/Swiss-Prot
Last modified
May 1, 2013.
Version 123.
History...
Names·Attributes·General annotation·Ontologies·Interactions·Alt products·Sequence annotation·Sequences·References·Cross-refs·Entry info·DocumentsCustomize order
Names·Attributes·General annotation·Ontologies·Interactions·Alt products·Sequence annotation·Sequences·References·Cross-refs·Entry info·DocumentsCustomize orderNames and origin
| Protein names | Recommended name: Exportin-2 Short name=Exp2 Alternative name(s): Cellular apoptosis susceptibility protein Chromosome segregation 1-like protein Importin-alpha re-exporter | ||||
| Gene names |
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| Organism | Homo sapiens (Human) [Reference proteome] | ||||
| Taxonomic identifier | 9606 [NCBI] | ||||
| Taxonomic lineage | Eukaryota › Metazoa › Chordata › Craniata › Vertebrata › Euteleostomi › Mammalia › Eutheria › Euarchontoglires › Primates › Haplorrhini › Catarrhini › Hominidae › Homo![]() |
Protein attributes
| Sequence length | 971 AA. |
| Sequence status | Complete. |
| Protein existence | Evidence at protein level |
General annotation (Comments)
| Function | Export receptor for importin-alpha. Mediates importin-alpha re-export from the nucleus to the cytoplasm after import substrates (cargos) have been released into the nucleoplasm. In the nucleus binds cooperatively to importin-alpha and to the GTPase Ran in its active GTP-bound form. Docking of this trimeric complex to the nuclear pore complex (NPC) is mediated through binding to nucleoporins. Upon transit of a nuclear export complex into the cytoplasm, disassembling of the complex and hydrolysis of Ran-GTP to Ran-GDP (induced by RANBP1 and RANGAP1, respectively) cause release of the importin-alpha from the export receptor. CSE1L/XPO2 then return to the nuclear compartment and mediate another round of transport. The directionality of nuclear export is thought to be conferred by an asymmetric distribution of the GTP- and GDP-bound forms of Ran between the cytoplasm and nucleus. Ref.8 |
| Subunit structure | Found in a complex with CSE1L/XPO2, Ran and KPNA2. Binds with high affinity to importin-alpha only in the presence of RanGTP. The complex is dissociated by the combined action of RanBP1 and RanGAP1. Ref.8 |
| Subcellular location | Cytoplasm. Nucleus. Note: Shuttles between the nucleus and the cytoplasm. |
| Tissue specificity | Highly expressed in proliferating cells. |
| Sequence similarities | Belongs to the XPO2/CSE1 family. Contains 1 importin N-terminal domain. |
Ontologies
| Keywords | |
|---|---|
| Biological process | Protein transport Transport |
| Cellular component | Cytoplasm Nucleus |
| Coding sequence diversity | Alternative splicing Polymorphism |
| PTM | Acetylation |
| Technical term | Complete proteome Direct protein sequencing Reference proteome |
| Gene Ontology (GO) | |
| Biological_process | apoptotic process Traceable author statement Ref.1. Source: ProtInc cell proliferationTraceable author statement Ref.1. Source: ProtInc |
| Cellular_component | cytoplasm Traceable author statement Ref.8. Source: ProtInc nuclear poreInferred from electronic annotation. Source: InterPro nucleusTraceable author statement Ref.8. Source: ProtInc |
| Molecular_function | importin-alpha export receptor activity Traceable author statement Ref.8. Source: ProtInc |
| Complete GO annotation... | |
Binary interactions
With | Entry | #Exp. | IntAct | Notes |
|---|---|---|---|---|
| TP53 | P04637 | 5 | EBI-286709,EBI-366083 |
Alternative products
| This entry describes 3 isoforms produced by alternative splicing. [Align] [Select] | ||||||
| Isoform 1 (identifier: P55060-1) Also known as: A; This isoform has been chosen as the 'canonical' sequence. All positional information in this entry refers to it. This is also the sequence that appears in the downloadable versions of the entry. | ||||||
| Isoform 2 (identifier: P55060-2) The sequence of this isoform differs from the canonical sequence as follows: 190-195: ATIELC → VWNASW 196-971: Missing. | ||||||
| Isoform 3 (identifier: P55060-3) The sequence of this isoform differs from the canonical sequence as follows: 943-945: VPS → TYF 946-971: Missing. |
Sequence annotation (Features)
| Feature key | Position(s) | Length | Description | Graphical view | Feature identifier | ||||
Molecule processing | |||||||||
|---|---|---|---|---|---|---|---|---|---|
| Chain | 1 – 971 | 971 | Exportin-2 | PRO_0000117287 | |||||
Regions | |||||||||
| Domain | 29 – 102 | 74 | Importin N-terminal | ||||||
Amino acid modifications | |||||||||
| Modified residue | 1 | 1 | N-acetylmethionine Ref.7 | ||||||
| Modified residue | 574 | 1 | N6-acetyllysine Ref.10 | ||||||
| Modified residue | 824 | 1 | N6-acetyllysine Ref.10 | ||||||
Natural variations | |||||||||
| Alternative sequence | 190 – 195 | 6 | ATIELC → VWNASW in isoform 2. | VSP_001222 | |||||
| Alternative sequence | 196 – 971 | 776 | Missing in isoform 2. | VSP_001223 | |||||
| Alternative sequence | 943 – 945 | 3 | VPS → TYF in isoform 3. | VSP_001224 | |||||
| Alternative sequence | 946 – 971 | 26 | Missing in isoform 3. | VSP_001225 | |||||
| Natural variant | 754 | 1 | I → V. Corresponds to variant rs2229042 [ dbSNP | Ensembl ]. | VAR_029327 | |||||
| Natural variant | 842 | 1 | C → F in a colorectal cancer sample; somatic mutation. Ref.12 | VAR_036558 | |||||
| Natural variant | 968 | 1 | V → L. Corresponds to variant rs3505 [ dbSNP | Ensembl ]. | VAR_048836 | |||||
Experimental info | |||||||||
| Sequence conflict | 231 – 233 | 3 | FED → WEG in AAC50367. Ref.1 | ||||||
| Sequence conflict | 231 – 233 | 3 | FED → WEG in AAC35008. Ref.2 | ||||||
| Sequence conflict | 514 | 1 | E → G in AAC50367. Ref.1 | ||||||
| Sequence conflict | 514 | 1 | E → G in AAC35008. Ref.2 | ||||||
| Sequence conflict | 848 | 1 | K → N in AAC50367. Ref.1 | ||||||
| Sequence conflict | 934 | 1 | K → M in AAC50367. Ref.1 | ||||||
Sequences
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References
| « Hide 'large scale' references | |
| [1] | "Cloning and characterization of a cellular apoptosis susceptibility gene, the human homologue to the yeast chromosome segregation gene CSE1." Brinkmann U., Brinkmann E., Gallo M., Pastan I. Proc. Natl. Acad. Sci. U.S.A. 92:10427-10431(1995) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 1). Tissue: Placenta. |
| [2] | "Tissue-specific alternative splicing of the CSE1L/CAS (cellular apoptosis susceptibility) gene." Brinkmann U., Brinkmann E., Bera T.K., Wellmann A., Pastan I. Genomics 58:41-49(1999) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [GENOMIC DNA / MRNA] (ISOFORMS 1; 2 AND 3). Tissue: Brain. |
| [3] | "Complete sequencing and characterization of 21,243 full-length human cDNAs." Ota T., Suzuki Y., Nishikawa T., Otsuki T., Sugiyama T., Irie R., Wakamatsu A., Hayashi K., Sato H., Nagai K., Kimura K., Makita H., Sekine M., Obayashi M., Nishi T., Shibahara T., Tanaka T., Ishii S. Sugano S.Nat. Genet. 36:40-45(2004) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM 1). Tissue: Cerebellum. |
| [4] | "The DNA sequence and comparative analysis of human chromosome 20." Deloukas P., Matthews L.H., Ashurst J.L., Burton J., Gilbert J.G.R., Jones M., Stavrides G., Almeida J.P., Babbage A.K., Bagguley C.L., Bailey J., Barlow K.F., Bates K.N., Beard L.M., Beare D.M., Beasley O.P., Bird C.P., Blakey S.E. Rogers J.Nature 414:865-871(2001) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]. |
| [5] | Mural R.J., Istrail S., Sutton G.G., Florea L., Halpern A.L., Mobarry C.M., Lippert R., Walenz B., Shatkay H., Dew I., Miller J.R., Flanigan M.J., Edwards N.J., Bolanos R., Fasulo D., Halldorsson B.V., Hannenhalli S., Turner R. Venter J.C.Submitted (SEP-2005) to the EMBL/GenBank/DDBJ databases Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]. |
| [6] | "The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC)." The MGC Project Team Genome Res. 14:2121-2127(2004) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM 1). Tissue: Testis. |
| [7] | Bienvenut W.V. Submitted (JUN-2005) to UniProtKB Cited for: PROTEIN SEQUENCE OF 1-16; 18-26; 32-67; 76-83; 94-109; 138-151; 166-217; 252-268; 282-288; 293-309; 332-371; 373-382; 396-418; 428-440; 446-481; 560-574; 698-736; 769-777; 789-816; 825-832 AND 913-934, ACETYLATION AT MET-1, MASS SPECTROMETRY. Tissue: B-cell lymphoma. |
| [8] | "Export of importin-alpha from the nucleus is mediated by a specific nuclear transport factor." Kutay U., Bischoff F.R., Kostka S., Kraft R., Goerlich D. Cell 90:1061-1071(1997) [PubMed] [Europe PMC] [Abstract] Cited for: PROTEIN SEQUENCE OF 356-370, FUNCTION IN PROTEIN NUCLEAR EXPORT, IDENTIFICATION IN A COMPLEX WITH RAN AND KPNA2. |
| [9] | "Determination of the functional domain organization of the importin alpha nuclear import factor." Herold A., Truant R., Wiegand H., Cullen B.R. J. Cell Biol. 143:309-318(1998) [PubMed] [Europe PMC] [Abstract] Cited for: INTERACTION WITH KPNA2. |
| [10] | "Lysine acetylation targets protein complexes and co-regulates major cellular functions." Choudhary C., Kumar C., Gnad F., Nielsen M.L., Rehman M., Walther T.C., Olsen J.V., Mann M. Science 325:834-840(2009) [PubMed] [Europe PMC] [Abstract] Cited for: ACETYLATION [LARGE SCALE ANALYSIS] AT LYS-574 AND LYS-824, MASS SPECTROMETRY. |
| [11] | "Initial characterization of the human central proteome." Burkard T.R., Planyavsky M., Kaupe I., Breitwieser F.P., Buerckstuemmer T., Bennett K.L., Superti-Furga G., Colinge J. BMC Syst. Biol. 5:17-17(2011) [PubMed] [Europe PMC] [Abstract] Cited for: IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]. |
| [12] | "The consensus coding sequences of human breast and colorectal cancers." Sjoeblom T., Jones S., Wood L.D., Parsons D.W., Lin J., Barber T.D., Mandelker D., Leary R.J., Ptak J., Silliman N., Szabo S., Buckhaults P., Farrell C., Meeh P., Markowitz S.D., Willis J., Dawson D., Willson J.K.V. Velculescu V.E.Science 314:268-274(2006) [PubMed] [Europe PMC] [Abstract] Cited for: VARIANT [LARGE SCALE ANALYSIS] PHE-842. |
| + | Additional computationally mapped references. |
Cross-references
Sequence databases | |
|---|---|
| EMBL GenBank DDBJ | U33286 mRNA. Translation: AAC50367.1. AF053640 mRNA. Translation: AAC35007.1. AF053641 mRNA. Translation: AAC35008.1. AF053642 mRNA. Translation: AAC35009.1. AF053651 AF053650 Genomic DNA. Translation: AAC35297.1.AK312306 mRNA. Translation: BAG35231.1. AL121903, AL133174 Genomic DNA. Translation: CAI19615.1. AL133174, AL121903 Genomic DNA. Translation: CAI42818.1. CH471077 Genomic DNA. Translation: EAW75676.1. CH471077 Genomic DNA. Translation: EAW75677.1. BC108309 mRNA. Translation: AAI08310.1. BC109313 mRNA. Translation: AAI09314.1. BC109314 mRNA. Translation: AAI09315.1. |
| IPI | IPI00022744. IPI00219762. IPI00219994. |
| PIR | I39166. |
| RefSeq | NP_001243064.1. NM_001256135.1. NP_001307.2. NM_001316.3. |
| UniGene | Hs.90073. |
3D structure databases | |
| ProteinModelPortal | P55060. |
| ModBase | Search... |
Protein-protein interaction databases | |
| IntAct | P55060. 19 interactions. |
| MINT | MINT-5001090. |
PTM databases | |
| PhosphoSite | P55060. |
Polymorphism databases | |
| DMDM | 62297557. |
Proteomic databases | |
| PaxDb | P55060. |
| PRIDE | P55060. |
Protocols and materials databases | |
| DNASU | 1434. |
| StructuralBiologyKnowledgebase | Search... |
Genome annotation databases | |
| Ensembl | ENST00000262982; ENSP00000262982; ENSG00000124207. |
| GeneID | 1434. |
| KEGG | hsa:1434. |
| UCSC | uc002xty.3. human. |
Organism-specific databases | |
| CTD | 1434. |
| GeneCards | GC20P047662. |
| HGNC | HGNC:2431. CSE1L. |
| HPA | CAB002140. |
| MIM | 601342. gene. |
| neXtProt | NX_P55060. |
| PharmGKB | PA26933. |
| GenAtlas | Search... |
Phylogenomic databases | |
| eggNOG | COG5657. |
| HOVERGEN | HBG080050. |
| InParanoid | P55060. |
| OMA | PEEYIRR. |
| OrthoDB | EOG4NGGKZ. |
| PhylomeDB | P55060. |
Gene expression databases | |
| ArrayExpress | P55060. |
| Bgee | P55060. |
| CleanEx | HS_CSE1L. |
| Genevestigator | P55060. |
| GermOnline | ENSG00000124207. Homo sapiens. |
Family and domain databases | |
| Gene3D | 1.25.10.10. 2 hits. |
| InterPro | IPR011989. ARM-like. IPR016024. ARM-type_fold. IPR005043. CAS_CSE1_C. IPR013713. Exportin/Importin_Cse1-like. IPR001494. Importin-beta_N. [Graphical view] |
| Pfam | PF03378. CAS_CSE1. 1 hit. PF08506. Cse1. 1 hit. PF03810. IBN_N. 1 hit. [Graphical view] |
| SMART | SM00913. IBN_N. 1 hit. [Graphical view] |
| SUPFAM | SSF48371. ARM-type_fold. 1 hit. |
| PROSITE | PS50166. IMPORTIN_B_NT. 1 hit. [Graphical view] |
| ProtoNet | Search... |
Other | |
| ChiTaRS | CSE1L. human. |
| GenomeRNAi | 1434. |
| NextBio | 5853. |
| SOURCE | Search... |
Entry information
| Entry name | XPO2_HUMAN | ||||||||
| Accession | Primary (citable) accession number: P55060 Secondary accession number(s): B2R5T4 Q9UPA0 | ||||||||
| Entry history |
| ||||||||
| Entry status | Reviewed (UniProtKB/Swiss-Prot) | ||||||||
| Annotation program | Chordata Protein Annotation Program | ||||||||
| Disclaimer | Any medical or genetic information present in this entry is provided for research, educational and informational purposes only. It is not in any way intended to be used as a substitute for professional medical advice, diagnosis, treatment or care. | ||||||||
Relevant documents
| Human chromosome 20 Human chromosome 20: entries, gene names and cross-references to MIM |
| Human entries with polymorphisms or disease mutations List of human entries with polymorphisms or disease mutations |
| Human polymorphisms and disease mutations Index of human polymorphisms and disease mutations |
| MIM cross-references Online Mendelian Inheritance in Man (MIM) cross-references in UniProtKB/Swiss-Prot |
| SIMILARITY comments Index of protein domains and families |

Clusters with
