Q8TEX9 (IPO4_HUMAN) Reviewed, UniProtKB/Swiss-Prot

Names and origin

Protein names

Recommended name:
Importin-4
Short name=Imp4

Alternative name(s):
Importin-4b
Short name=Imp4b
Ran-binding protein 4
Short name=RanBP4

Gene names

Name:	IPO4
Synonyms:	IMP4B, RANBP4

Organism

Homo sapiens (Human) [Reference proteome]

Taxonomic identifier

9606 [NCBI]

Taxonomic lineage

cellular organisms › Eukaryota › Opisthokonta › Metazoa › Eumetazoa › Bilateria › Deuterostomia › Chordata › Craniata › Vertebrata › Gnathostomata › Teleostomi › Euteleostomi › Sarcopterygii › Tetrapoda › Amniota › Mammalia › Theria › Eutheria › Euarchontoglires › Primates › Haplorrhini › Simiiformes › Catarrhini › Hominoidea › Hominidae › Homininae › Homo

Protein attributes

Sequence length	1081 AA.
Sequence status	Complete.
Protein existence	Evidence at protein level

General annotation (Comments)

Function	Functions in nuclear protein import as nuclear transport receptor. Serves as receptor for nuclear localization signals (NLS) in cargo substrates. Is thought to mediate docking of the importin/substrate complex to the nuclear pore complex (NPC) through binding to nucleoporin and the complex is subsequently translocated through the pore by an energy requiring, Ran-dependent mechanism. At the nucleoplasmic side of the NPC, Ran binds to the importin, the importin/substrate complex dissociates and importin is re-exported from the nucleus to the cytoplasm where GTP hydrolysis releases Ran. The directionality of nuclear import is thought to be conferred by an asymmetric distribution of the GTP- and GDP-bound forms of Ran between the cytoplasm and nucleus By similarity. Mediates the nuclear import of RPS3A. In vitro, mediates the nuclear import of human cytomegalovirus UL84 by recognizing a non-classical NLS. Ref.1
Subunit structure	Binds with high affinity to RPS3A. The binding is coupled to RanGTP cycles. Binds to human cytomegalovirus UL84. Found in a cytosolic complex with ASF1A, ASF1B, CDAN1 and histones H3.1 and H4. Ref.1 Ref.8 Ref.10
Subcellular location	Cytoplasm By similarity. Nucleus By similarity.
Sequence similarities	Belongs to the importin beta family. Contains 6 HEAT repeats. Contains 1 importin N-terminal domain.
Sequence caution	The sequence BAB15616.1 differs from that shown. Reason: Frameshift at position 234. The sequence BAB55421.1 differs from that shown. Reason: Erroneous initiation. Translation N-terminally shortened. The sequence BAC11174.1 differs from that shown. Reason: Erroneous initiation. Translation N-terminally extended. The sequence CAD62595.1 differs from that shown. Reason: Erroneous initiation. Translation N-terminally extended.

Ontologies

Keywords
Biological process	Protein transport Transport
Cellular component	Cytoplasm Nucleus
Coding sequence diversity	Alternative splicing Polymorphism
Domain	Repeat
PTM	Acetylation
Technical term	Complete proteome Direct protein sequencing Reference proteome
Gene Ontology (GO)
Biological_process	intracellular protein transport Inferred from electronic annotation. Source: InterPro protein transport Non-traceable author statement Ref.1. Source: UniProtKB
Cellular_component	cytoplasm Inferred from electronic annotation. Source: UniProtKB-SubCell nuclear pore Inferred from electronic annotation. Source: InterPro
Molecular_function	protein transporter activity Inferred from electronic annotation. Source: InterPro
Complete GO annotation...

Alternative products

This entry describes 2 isoforms produced by alternative splicing. [Align] [Select]

Isoform 1 (identifier: Q8TEX9-1)

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: Q8TEX9-2)

The sequence of this isoform differs from the canonical sequence as follows:
     1038-1038: P → PGE

Note: No experimental confirmation available.

Sequence annotation (Features)

	Feature key	Position(s)	Length	Description	Graphical view	Feature identifier
Molecule processing
	Chain	1 – 1081	1081	Importin-4		PRO_0000120748
Regions
	Domain	24 – 90	67	Importin N-terminal
	Repeat	348 – 385	38	HEAT 1
	Repeat	390 – 427	38	HEAT 2
	Repeat	431 – 471	41	HEAT 3
	Repeat	475 – 513	39	HEAT 4
	Repeat	895 – 932	38	HEAT 5
	Repeat	936 – 974	39	HEAT 6
Amino acid modifications
	Modified residue	1	1	N-acetylmethionine Ref.6
Natural variations
	Alternative sequence	1038	1	P → PGE in isoform 2.		VSP_009339
	Natural variant	513	1	A → V. Ref.1 Ref.7 Corresponds to variant rs7146310 [ dbSNP | Ensembl ].		VAR_030758
	Natural variant	580	1	P → A. Ref.1 Ref.7 Corresponds to variant rs11550452 [ dbSNP | Ensembl ].		VAR_030759
Experimental info
	Sequence conflict	865	1	K → R in BAB15616. Ref.7
	Sequence conflict	943	1	G → E in BAB55421. Ref.7

Sequences

	Sequence		Length	Mass (Da)
	Isoform 1 [UniParc]. Last modified February 20, 2007. Version 2. Checksum: 36D11AF0D6DA8B1E	FASTA	1,081	118,715
	10 20 30 40 50 60 MESAGLEQLL RELLLPDTER IRRATEQLQI VLRAPAALPA LCDLLASAAD PQIRQFAAVL 70 80 90 100 110 120 TRRRLNTRWR RLAAEQRESL KSLILTALQR ETEHCVSLSL AQLSATIFRK EGLEAWPQLL 130 140 150 160 170 180 QLLQHSTHSP HSPEREMGLL LLSVVVTSRP EAFQPHHREL LRLLNETLGE VGSPGLLFYS 190 200 210 220 230 240 LRTLTTMAPY LSTEDVPLAR MLVPKLIMAM QTLIPIDEAK ACEALEALDE LLESEVPVIT 250 260 270 280 290 300 PYLSEVLTFC LEVARNVALG NAIRIRILCC LTFLVKVKSK ALLKNRLLPP LLHTLFPIVA 310 320 330 340 350 360 AEPPPGQLDP EDQDSEEEEL EIELMGETPK HFAVQVVDML ALHLPPEKLC PQLMPMLEEA 370 380 390 400 410 420 LRSESPYQRK AGLLVLAVLS DGAGDHIRQR LLPPLLQIVC KGLEDPSQVV RNAALFALGQ 430 440 450 460 470 480 FSENLQPHIS SYSREVMPLL LAYLKSVPLG HTHHLAKACY ALENFVENLG PKVQPYLPEL 490 500 510 520 530 540 MECMLQLLRN PSSPRAKELA VSALGAIATA AQASLLPYFP AIMEHLREFL LTGREDLQPV 550 560 570 580 590 600 QIQSLETLGV LARAVGEPMR PLAEECCQLG LGLCDQVDDP DLRRCTYSLF AALSGLMGEG 610 620 630 640 650 660 LAPHLEQITT LMLLSLRSTE GIVPQYDGSS SFLLFDDESD GEEEEELMDE DVEEEDDSEI 670 680 690 700 710 720 SGYSVENAFF DEKEDTCAAV GEISVNTSVA FLPYMESVFE EVFKLLECPH LNVRKAAHEA 730 740 750 760 770 780 LGQFCCALHK ACQSCPSEPN TAALQAALAR VVPSYMQAVN RERERQVVMA VLEALTGVLR 790 800 810 820 830 840 SCGTLTLKPP GRLAELCGVL KAVLQRKTAC QDTDEEEEEE DDDQAEYDAM LLEHAGEAIP 850 860 870 880 890 900 ALAAAAGGDS FAPFFAGFLP LLVCKTKQGC TVAEKSFAVG TLAETIQGLG AASAQFVSRL 910 920 930 940 950 960 LPVLLSTAQE ADPEVRSNAI FGMGVLAEHG GHPAQEHFPK LLGLLFPLLA RERHDRVRDN 970 980 990 1000 1010 1020 ICGALARLLM ASPTRKPEPQ VLAALLHALP LKEDLEEWVT IGRLFSFLYQ SSPDQVIDVA 1030 1040 1050 1060 1070 1080 PELLRICSLI LADNKIPPDT KAALLLLLTF LAKQHTDSFQ AALGSLPVDK AQELQAVLGL S
	Isoform 2 [UniParc]. Checksum: BBB539F050812D96	FASTA	1,083	118,901
	10 20 30 40 50 60 MESAGLEQLL RELLLPDTER IRRATEQLQI VLRAPAALPA LCDLLASAAD PQIRQFAAVL 70 80 90 100 110 120 TRRRLNTRWR RLAAEQRESL KSLILTALQR ETEHCVSLSL AQLSATIFRK EGLEAWPQLL 130 140 150 160 170 180 QLLQHSTHSP HSPEREMGLL LLSVVVTSRP EAFQPHHREL LRLLNETLGE VGSPGLLFYS 190 200 210 220 230 240 LRTLTTMAPY LSTEDVPLAR MLVPKLIMAM QTLIPIDEAK ACEALEALDE LLESEVPVIT 250 260 270 280 290 300 PYLSEVLTFC LEVARNVALG NAIRIRILCC LTFLVKVKSK ALLKNRLLPP LLHTLFPIVA 310 320 330 340 350 360 AEPPPGQLDP EDQDSEEEEL EIELMGETPK HFAVQVVDML ALHLPPEKLC PQLMPMLEEA 370 380 390 400 410 420 LRSESPYQRK AGLLVLAVLS DGAGDHIRQR LLPPLLQIVC KGLEDPSQVV RNAALFALGQ 430 440 450 460 470 480 FSENLQPHIS SYSREVMPLL LAYLKSVPLG HTHHLAKACY ALENFVENLG PKVQPYLPEL 490 500 510 520 530 540 MECMLQLLRN PSSPRAKELA VSALGAIATA AQASLLPYFP AIMEHLREFL LTGREDLQPV 550 560 570 580 590 600 QIQSLETLGV LARAVGEPMR PLAEECCQLG LGLCDQVDDP DLRRCTYSLF AALSGLMGEG 610 620 630 640 650 660 LAPHLEQITT LMLLSLRSTE GIVPQYDGSS SFLLFDDESD GEEEEELMDE DVEEEDDSEI 670 680 690 700 710 720 SGYSVENAFF DEKEDTCAAV GEISVNTSVA FLPYMESVFE EVFKLLECPH LNVRKAAHEA 730 740 750 760 770 780 LGQFCCALHK ACQSCPSEPN TAALQAALAR VVPSYMQAVN RERERQVVMA VLEALTGVLR 790 800 810 820 830 840 SCGTLTLKPP GRLAELCGVL KAVLQRKTAC QDTDEEEEEE DDDQAEYDAM LLEHAGEAIP 850 860 870 880 890 900 ALAAAAGGDS FAPFFAGFLP LLVCKTKQGC TVAEKSFAVG TLAETIQGLG AASAQFVSRL 910 920 930 940 950 960 LPVLLSTAQE ADPEVRSNAI FGMGVLAEHG GHPAQEHFPK LLGLLFPLLA RERHDRVRDN 970 980 990 1000 1010 1020 ICGALARLLM ASPTRKPEPQ VLAALLHALP LKEDLEEWVT IGRLFSFLYQ SSPDQVIDVA 1030 1040 1050 1060 1070 1080 PELLRICSLI LADNKIPPGE DTKAALLLLL TFLAKQHTDS FQAALGSLPV DKAQELQAVL GLS

References

	« Hide 'large scale' referencesShow 'large scale' references »
[1]	"Importins fulfill a dual function as nuclear import receptors and cytoplasmic chaperones for exposed basic domains." Jaekel S., Mingot J.-M., Schwarzmaier P., Hartmann E., Goerlich D. EMBO J. 21:377-386(2002) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 1), FUNCTION, INTERACTION WITH RPS3A, VARIANTS VAL-513 AND ALA-580.
[2]	"Full-length cDNA libraries and normalization." Li W.B., Gruber C., Jessee J., Polayes D. Submitted (FEB-2003) to the EMBL/GenBank/DDBJ databases Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM 1). Tissue: Neuroblastoma.
[3]	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. , Yooseph S., Lu F., Nusskern D.R., Shue B.C., Zheng X.H., Zhong F., Delcher A.L., Huson D.H., Kravitz S.A., Mouchard L., Reinert K., Remington K.A., Clark A.G., Waterman M.S., Eichler E.E., Adams M.D., Hunkapiller M.W., Myers E.W., Venter J.C. Submitted (SEP-2005) to the EMBL/GenBank/DDBJ databases Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
[4]	"The DNA sequence and analysis of human chromosome 14." Heilig R., Eckenberg R., Petit J.-L., Fonknechten N., Da Silva C., Cattolico L., Levy M., Barbe V., De Berardinis V., Ureta-Vidal A., Pelletier E., Vico V., Anthouard V., Rowen L., Madan A., Qin S., Sun H., Du H. , Pepin K., Artiguenave F., Robert C., Cruaud C., Bruels T., Jaillon O., Friedlander L., Samson G., Brottier P., Cure S., Segurens B., Aniere F., Samain S., Crespeau H., Abbasi N., Aiach N., Boscus D., Dickhoff R., Dors M., Dubois I., Friedman C., Gouyvenoux M., James R., Madan A., Mairey-Estrada B., Mangenot S., Martins N., Menard M., Oztas S., Ratcliffe A., Shaffer T., Trask B., Vacherie B., Bellemere C., Belser C., Besnard-Gonnet M., Bartol-Mavel D., Boutard M., Briez-Silla S., Combette S., Dufosse-Laurent V., Ferron C., Lechaplais C., Louesse C., Muselet D., Magdelenat G., Pateau E., Petit E., Sirvain-Trukniewicz P., Trybou A., Vega-Czarny N., Bataille E., Bluet E., Bordelais I., Dubois M., Dumont C., Guerin T., Haffray S., Hammadi R., Muanga J., Pellouin V., Robert D., Wunderle E., Gauguet G., Roy A., Sainte-Marthe L., Verdier J., Verdier-Discala C., Hillier L.W., Fulton L., McPherson J., Matsuda F., Wilson R., Scarpelli C., Gyapay G., Wincker P., Saurin W., Quetier F., Waterston R., Hood L., Weissenbach J. Nature 421:601-607(2003) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
[5]	"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: Brain, Placenta and Skin.
[6]	Bienvenut W.V. Submitted (JUN-2005) to UniProtKB Cited for: PROTEIN SEQUENCE OF 1-11; 24-33; 55-62; 82-90; 402-411; 528-534; 751-761; 900-916; 941-951 AND 1042-1053, ACETYLATION AT MET-1, MASS SPECTROMETRY. Tissue: B-cell lymphoma.
[7]	"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. , Yamamoto J., Saito K., Kawai Y., Isono Y., Nakamura Y., Nagahari K., Murakami K., Yasuda T., Iwayanagi T., Wagatsuma M., Shiratori A., Sudo H., Hosoiri T., Kaku Y., Kodaira H., Kondo H., Sugawara M., Takahashi M., Kanda K., Yokoi T., Furuya T., Kikkawa E., Omura Y., Abe K., Kamihara K., Katsuta N., Sato K., Tanikawa M., Yamazaki M., Ninomiya K., Ishibashi T., Yamashita H., Murakawa K., Fujimori K., Tanai H., Kimata M., Watanabe M., Hiraoka S., Chiba Y., Ishida S., Ono Y., Takiguchi S., Watanabe S., Yosida M., Hotuta T., Kusano J., Kanehori K., Takahashi-Fujii A., Hara H., Tanase T.-O., Nomura Y., Togiya S., Komai F., Hara R., Takeuchi K., Arita M., Imose N., Musashino K., Yuuki H., Oshima A., Sasaki N., Aotsuka S., Yoshikawa Y., Matsunawa H., Ichihara T., Shiohata N., Sano S., Moriya S., Momiyama H., Satoh N., Takami S., Terashima Y., Suzuki O., Nakagawa S., Senoh A., Mizoguchi H., Goto Y., Shimizu F., Wakebe H., Hishigaki H., Watanabe T., Sugiyama A., Takemoto M., Kawakami B., Yamazaki M., Watanabe K., Kumagai A., Itakura S., Fukuzumi Y., Fujimori Y., Komiyama M., Tashiro H., Tanigami A., Fujiwara T., Ono T., Yamada K., Fujii Y., Ozaki K., Hirao M., Ohmori Y., Kawabata A., Hikiji T., Kobatake N., Inagaki H., Ikema Y., Okamoto S., Okitani R., Kawakami T., Noguchi S., Itoh T., Shigeta K., Senba T., Matsumura K., Nakajima Y., Mizuno T., Morinaga M., Sasaki M., Togashi T., Oyama M., Hata H., Watanabe M., Komatsu T., Mizushima-Sugano J., Satoh T., Shirai Y., Takahashi Y., Nakagawa K., Okumura K., Nagase T., Nomura N., Kikuchi H., Masuho Y., Yamashita R., Nakai K., Yada T., Nakamura Y., Ohara O., Isogai T., Sugano S. Nat. Genet. 36:40-45(2004) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] OF 233-1081 (ISOFORM 1), NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] OF 538-1081 (ISOFORM 2), VARIANTS VAL-513 AND ALA-580.
[8]	"A nonconventional nuclear localization signal within the UL84 protein of human cytomegalovirus mediates nuclear import via the importin alpha/beta pathway." Lischka P., Sorg G., Kann M., Winkler M., Stamminger T. J. Virol. 77:3734-3748(2003) [PubMed] [Europe PMC] [Abstract] Cited for: INTERACTION WITH HCMV UL84.
[9]	"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].
[10]	"Codanin-1, mutated in the anaemic disease CDAI, regulates Asf1 function in S-phase histone supply." Ask K., Jasencakova Z., Menard P., Feng Y., Almouzni G., Groth A. EMBO J. 31:2013-2023(2012) [PubMed] [Europe PMC] [Abstract] Cited for: IDENTIFICATION IN A COMPLEX WITH ASF1A; ASF1B; CDNA1; HISTONES H3.2 AND H4.
+	Additional computationally mapped references.

Cross-references

Sequence databases
EMBL GenBank DDBJ	AF411122 mRNA. Translation: AAL78660.1. BX248267 mRNA. Translation: CAD62595.1. Different initiation. AL136295 Genomic DNA. No translation available. CH471078 Genomic DNA. Translation: EAW66078.1. BC003690 mRNA. Translation: AAH03690.2. BC110804 mRNA. Translation: AAI10805.1. BC136759 mRNA. Translation: AAI36760.1. AK026991 mRNA. Translation: BAB15616.1. Frameshift. AK027871 mRNA. Translation: BAB55421.1. Different initiation. AK074743 mRNA. Translation: BAC11174.1. Different initiation.
IPI	IPI00156374. IPI00398009.
RefSeq	NP_078934.3. NM_024658.3.
UniGene	Hs.411865.
3D structure databases
ProteinModelPortal	Q8TEX9.
ModBase	Search...
Protein-protein interaction databases
DIP	DIP-32944N.
IntAct	Q8TEX9. 8 interactions.
STRING	9606.ENSP00000346453.
PTM databases
PhosphoSite	Q8TEX9.
Polymorphism databases
DMDM	126302558.
Proteomic databases
PaxDb	Q8TEX9.
PRIDE	Q8TEX9.
Protocols and materials databases
StructuralBiologyKnowledgebase	Search...
Genome annotation databases
Ensembl	ENST00000354464; ENSP00000346453; ENSG00000196497.
GeneID	79711.
KEGG	hsa:79711.
UCSC	uc001wmt.1. human. uc001wmu.2. human.
Organism-specific databases
CTD	79711.
GeneCards	GC14M024649.
HGNC	HGNC:19426. IPO4.
HPA	HPA039043.
neXtProt	NX_Q8TEX9.
PharmGKB	PA134968932.
GenAtlas	Search...
Phylogenomic databases
eggNOG	NOG249123.
HOVERGEN	HBG049052.
InParanoid	Q8TEX9.
OMA	QRETEHS.
OrthoDB	EOG48GW2M.
Gene expression databases
ArrayExpress	Q8TEX9.
Bgee	Q8TEX9.
CleanEx	HS_IPO4.
Genevestigator	Q8TEX9.
GermOnline	ENSG00000196497. Homo sapiens.
Family and domain databases
Gene3D	1.25.10.10. 3 hits.
InterPro	IPR011989. ARM-like. IPR016024. ARM-type_fold. IPR021133. HEAT_type_2. IPR001494. Importin-beta_N. [Graphical view]
Pfam	PF03810. IBN_N. 1 hit. [Graphical view]
SMART	SM00913. IBN_N. 1 hit. [Graphical view]
SUPFAM	SSF48371. ARM-type_fold. 2 hits.
PROSITE	PS50077. HEAT_REPEAT. 2 hits. PS50166. IMPORTIN_B_NT. 1 hit. [Graphical view]
ProtoNet	Search...
Other
ChiTaRS	IPO4. human.
GenomeRNAi	79711.
NextBio	69037.

Entry information

Entry name

IPO4_HUMAN

Accession

Primary (citable) accession number: Q8TEX9
Secondary accession number(s): B2RN95 , Q2NL96, Q86TZ9, Q8NCG8, Q96SJ3, Q9BTI4, Q9H5L0

Entry history

Integrated into UniProtKB/Swiss-Prot:	February 2, 2004
Last sequence update:	February 20, 2007
Last modified:	May 1, 2013
This is version 112 of the entry and version 2 of the sequence. [Complete 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.