Q96P70 (IPO9_HUMAN) Reviewed, UniProtKB/Swiss-Prot

Names and origin

Protein names

Recommended name:
Importin-9
Short name=Imp9

Alternative name(s):
Ran-binding protein 9
Short name=RanBP9

Gene names

Name:	IPO9
Synonyms:	IMP9, KIAA1192, RANBP9
ORF Names:	HSPC273

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	1041 AA.
Sequence status	Complete.
Sequence processing	The displayed sequence is further processed into a mature form.
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 H2B histone By similarity, RPS7 and RPL18A. Prevents the cytoplasmic aggregation of RPS7 and RPL18A by shielding exposed basic domains. May also import H2A, H3, H4 histones By similarity, RPL4 and RPL6. Ref.1
Subunit structure	Binds with high affinity to RPS7 and RPL18A. The binding is coupled to RanGTP cycles. May bind H2A, H3, H4 histones By similarity, RPL4 and RPL6 with low affinity. Interacts with PPP2R1A and PPP2R1B. Ref.1 Ref.10
Subcellular location	Cytoplasm By similarity. Nucleus By similarity.
Sequence similarities	Belongs to the importin beta family. Contains 1 importin N-terminal domain.
Sequence caution	The sequence AAF28951.1 differs from that shown. Reason: Frameshift at position 982. The sequence BAA86506.1 differs from that shown. Reason: Erroneous initiation. The sequence BAA91588.1 differs from that shown. Reason: Erroneous initiation. The sequence BAB55181.1 differs from that shown. Reason: Erroneous initiation. The sequence BAC11173.1 differs from that shown. Reason: Erroneous initiation.

Ontologies

Keywords
Biological process	Protein transport Transport
Cellular component	Cytoplasm Nucleus
PTM	Acetylation
Technical term	Complete proteome Direct protein sequencing Reference proteome
Gene Ontology (GO)
Biological_process	protein import into nucleus Non-traceable author statement Ref.1. Source: UniProtKB
Cellular_component	cytoplasm Inferred from direct assay Ref.1. Source: UniProtKB nuclear pore Inferred from electronic annotation. Source: InterPro
Molecular_function	histone binding Inferred from sequence or structural similarity. Source: UniProtKB protein transporter activity Inferred from direct assay Ref.1. Source: UniProtKB
Complete GO annotation...

Sequence annotation (Features)

	Feature key	Position(s)	Length	Description	Graphical view	Feature identifier
Molecule processing
	Initiator methionine	1	1	Removed Ref.5
	Chain	2 – 1041	1040	Importin-9		PRO_0000120754
Regions
	Domain	43 – 119	77	Importin N-terminal
Amino acid modifications
	Modified residue	2	1	N-acetylalanine Ref.5
Experimental info
	Sequence conflict	632	1	E → G in BAC11173. Ref.4
	Sequence conflict	916	1	K → R in BAC11173. Ref.4
	Sequence conflict	935	1	R → P in BAB55181. Ref.4
	Sequence conflict	960	1	E → G in BAC11173. Ref.4

Sequences

	Sequence		Length	Mass (Da)
	Q96P70 [UniParc]. Last modified January 23, 2007. Version 3. Checksum: A1842C357AEDFD90	FASTA	1,041	115,963
	10 20 30 40 50 60 MAAAAAAGAA SGLPGPVAQG LKEALVDTLT GILSPVQEVR AAAEEQIKVL EVTEEFGVHL 70 80 90 100 110 120 AELTVDPQGA LAIRQLASVI LKQYVETHWC AQSEKFRPPE TTERAKIVIR ELLPNGLRES 130 140 150 160 170 180 ISKVRSSVAY AVSAIAHWDW PEAWPQLFNL LMEMLVSGDL NAVHGAMRVL TEFTREVTDT 190 200 210 220 230 240 QMPLVAPVIL PEMYKIFTMA EVYGIRTRSR AVEIFTTCAH MICNMEELEK GAAKVLIFPV 250 260 270 280 290 300 VQQFTEAFVQ ALQIPDGPTS DSGFKMEVLK AVTALVKNFP KHMVSSMQQI LPIVWNTLTE 310 320 330 340 350 360 SAAFYVRTEV NYTEEVEDPV DSDGEVLGFE NLVFSIFEFV HALLENSKFK STVKKALPEL 370 380 390 400 410 420 IYYIILYMQI TEEQIKVWTA NPQQFVEDED DDTFSYTVRI AAQDLLLAVA TDFQNESAAA 430 440 450 460 470 480 LAAAATRHLQ EAEQTKNSGT EHWWKIHEAC MLALGSVKAI ITDSVKNGRI HFDMHGFLTN 490 500 510 520 530 540 VILADLNLSV SPFLLGRALW AASRFTVAMS PELIQQFLQA TVSGLHETQP PSVRISAVRA 550 560 570 580 590 600 IWGYCDQLKV SESTHVLQPF LPSILDGLIH LAAQFSSEVL NLVMETLCIV CTVDPEFTAS 610 620 630 640 650 660 MESKICPFTI AIFLKYSNDP VVASLAQDIF KELSQIEACQ GPMQMRLIPT LVSIMQAPAD 670 680 690 700 710 720 KIPAGLCATA IDILTTVVRN TKPPLSQLLI CQAFPAVAQC TLHTDDNATM QNGGECLRAY 730 740 750 760 770 780 VSVTLEQVAQ WHDEQGHNGL WYVMQVVSQL LDPRTSEFTA AFVGRLVSTL ISKAGRELGE 790 800 810 820 830 840 NLDQILRAIL SKMQQAETLS VMQSLIMVFA HLVHTQLEPL LEFLCSLPGP TGKPALEFVM 850 860 870 880 890 900 AEWTSRQHLF YGQYEGKVSS VALCKLLQHG INADDKRLQD IRVKGEEIYS MDEGIRTRSK 910 920 930 940 950 960 SAKNPERWTN IPLLVKILKL IINELSNVME ANAARQATPA EWSQDDSNDM WEDQEEEEEE 970 980 990 1000 1010 1020 EEDGLAGQLL SDILATSKYE EDYYEDDEED DPDALKDPLY QIDLQAYLTD FLCQFAQQPC 1030 1040 YIMFSGHLND NERRVLQTIG I

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], FUNCTION, INTERACTION WITH RPS7; RPL18A; RPL4 AND RPL6.
[2]	"The DNA sequence and biological annotation of human chromosome 1." Gregory S.G., Barlow K.F., McLay K.E., Kaul R., Swarbreck D., Dunham A., Scott C.E., Howe K.L., Woodfine K., Spencer C.C.A., Jones M.C., Gillson C., Searle S., Zhou Y., Kokocinski F., McDonald L., Evans R., Phillips K. , Atkinson A., Cooper R., Jones C., Hall R.E., Andrews T.D., Lloyd C., Ainscough R., Almeida J.P., Ambrose K.D., Anderson F., Andrew R.W., Ashwell R.I.S., Aubin K., Babbage A.K., Bagguley C.L., Bailey J., Beasley H., Bethel G., Bird C.P., Bray-Allen S., Brown J.Y., Brown A.J., Buckley D., Burton J., Bye J., Carder C., Chapman J.C., Clark S.Y., Clarke G., Clee C., Cobley V., Collier R.E., Corby N., Coville G.J., Davies J., Deadman R., Dunn M., Earthrowl M., Ellington A.G., Errington H., Frankish A., Frankland J., French L., Garner P., Garnett J., Gay L., Ghori M.R.J., Gibson R., Gilby L.M., Gillett W., Glithero R.J., Grafham D.V., Griffiths C., Griffiths-Jones S., Grocock R., Hammond S., Harrison E.S.I., Hart E., Haugen E., Heath P.D., Holmes S., Holt K., Howden P.J., Hunt A.R., Hunt S.E., Hunter G., Isherwood J., James R., Johnson C., Johnson D., Joy A., Kay M., Kershaw J.K., Kibukawa M., Kimberley A.M., King A., Knights A.J., Lad H., Laird G., Lawlor S., Leongamornlert D.A., Lloyd D.M., Loveland J., Lovell J., Lush M.J., Lyne R., Martin S., Mashreghi-Mohammadi M., Matthews L., Matthews N.S.W., McLaren S., Milne S., Mistry S., Moore M.J.F., Nickerson T., O'Dell C.N., Oliver K., Palmeiri A., Palmer S.A., Parker A., Patel D., Pearce A.V., Peck A.I., Pelan S., Phelps K., Phillimore B.J., Plumb R., Rajan J., Raymond C., Rouse G., Saenphimmachak C., Sehra H.K., Sheridan E., Shownkeen R., Sims S., Skuce C.D., Smith M., Steward C., Subramanian S., Sycamore N., Tracey A., Tromans A., Van Helmond Z., Wall M., Wallis J.M., White S., Whitehead S.L., Wilkinson J.E., Willey D.L., Williams H., Wilming L., Wray P.W., Wu Z., Coulson A., Vaudin M., Sulston J.E., Durbin R.M., Hubbard T., Wooster R., Dunham I., Carter N.P., McVean G., Ross M.T., Harrow J., Olson M.V., Beck S., Rogers J., Bentley D.R. Nature 441:315-321(2006) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
[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 (JUL-2005) to the EMBL/GenBank/DDBJ databases Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
[4]	"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 1-276 AND 358-1041. Tissue: Amygdala.
[5]	Bienvenut W.V. Submitted (JUN-2005) to UniProtKB Cited for: PROTEIN SEQUENCE OF 2-22; 400-427 AND 908-916, CLEAVAGE OF INITIATOR METHIONINE, ACETYLATION AT ALA-2, MASS SPECTROMETRY. Tissue: B-cell lymphoma.
[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] OF 406-1041. Tissue: Placenta.
[7]	"The full-ORF clone resource of the German cDNA consortium." Bechtel S., Rosenfelder H., Duda A., Schmidt C.P., Ernst U., Wellenreuther R., Mehrle A., Schuster C., Bahr A., Bloecker H., Heubner D., Hoerlein A., Michel G., Wedler H., Koehrer K., Ottenwaelder B., Poustka A., Wiemann S., Schupp I. BMC Genomics 8:399-399(2007) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] OF 632-1041. Tissue: Amygdala.
[8]	"Cloning and functional analysis of cDNAs with open reading frames for 300 previously undefined genes expressed in CD34+ hematopoietic stem/progenitor cells." Zhang Q.-H., Ye M., Wu X.-Y., Ren S.-X., Zhao M., Zhao C.-J., Fu G., Shen Y., Fan H.-Y., Lu G., Zhong M., Xu X.-R., Han Z.-G., Zhang J.-W., Tao J., Huang Q.-H., Zhou J., Hu G.-X. , Gu J., Chen S.-J., Chen Z. Genome Res. 10:1546-1560(2000) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] OF 767-1041. Tissue: Umbilical cord blood.
[9]	"Characterization of cDNA clones selected by the GeneMark analysis from size-fractionated cDNA libraries from human brain." Hirosawa M., Nagase T., Ishikawa K., Kikuno R., Nomura N., Ohara O. DNA Res. 6:329-336(1999) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] OF 780-1041. Tissue: Brain.
[10]	"Interaction between protein phosphatase 2A and members of the importin beta superfamily." Lubert E.J., Sarge K.D. Biochem. Biophys. Res. Commun. 303:908-913(2003) [PubMed] [Europe PMC] [Abstract] Cited for: INTERACTION WITH PPP2R1A AND PPP2R1B.
[11]	"Kinase-selective enrichment enables quantitative phosphoproteomics of the kinome across the cell cycle." Daub H., Olsen J.V., Bairlein M., Gnad F., Oppermann F.S., Korner R., Greff Z., Keri G., Stemmann O., Mann M. Mol. Cell 31:438-448(2008) [PubMed] [Europe PMC] [Abstract] Cited for: IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS]. Tissue: Cervix carcinoma.
[12]	"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].
+	Additional computationally mapped references.

Cross-references

Sequence databases
EMBL GenBank DDBJ	AF410465 mRNA. Translation: AAL01416.1. AL645504 Genomic DNA. Translation: CAI17015.1. AK001264 mRNA. Translation: BAA91588.1. Different initiation. AK027532 mRNA. Translation: BAB55181.1. Different initiation. AK074740 mRNA. Translation: BAC11173.1. Different initiation. AK094603 mRNA. Translation: BAC04383.1. Sequence problems. CH471067 Genomic DNA. Translation: EAW91376.1. BC003604 mRNA. Translation: AAH03604.2. AF161391 mRNA. Translation: AAF28951.1. Frameshift. AL834323 mRNA. Translation: CAD38991.1. AB033018 mRNA. Translation: BAA86506.1. Different initiation.
IPI	IPI00185146.
RefSeq	NP_060555.2. NM_018085.4.
UniGene	Hs.596014.
3D structure databases
ProteinModelPortal	Q96P70.
ModBase	Search...
Protein-protein interaction databases
IntAct	Q96P70. 4 interactions.
MINT	MINT-1153224.
PTM databases
PhosphoSite	Q96P70.
Polymorphism databases
DMDM	41688593.
Proteomic databases
PaxDb	Q96P70.
PeptideAtlas	Q96P70.
PRIDE	Q96P70.
Protocols and materials databases
StructuralBiologyKnowledgebase	Search...
Genome annotation databases
Ensembl	ENST00000361565; ENSP00000354742; ENSG00000198700.
GeneID	55705.
KEGG	hsa:55705.
UCSC	uc001gwz.3. human.
Organism-specific databases
CTD	55705.
GeneCards	GC01P201798.
HGNC	HGNC:19425. IPO9.
neXtProt	NX_Q96P70.
PharmGKB	PA134930111.
HUGE	Search...
GenAtlas	Search...
Phylogenomic databases
eggNOG	COG5657.
HOVERGEN	HBG049054.
InParanoid	Q96P70.
OMA	ALQMPDG.
OrthoDB	EOG4NZTSH.
Gene expression databases
ArrayExpress	Q96P70.
Bgee	Q96P70.
CleanEx	HS_IPO9. HS_RANBP9.
Genevestigator	Q96P70.
GermOnline	ENSG00000198700. Homo sapiens.
Family and domain databases
Gene3D	1.25.10.10. 3 hits.
InterPro	IPR011989. ARM-like. IPR016024. ARM-type_fold. 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. 1 hit.
PROSITE	PS50166. IMPORTIN_B_NT. 1 hit. [Graphical view]
ProtoNet	Search...
Other
ChiTaRS	IPO9. human.
GenomeRNAi	55705.
NextBio	60559.

Entry information

Entry name

IPO9_HUMAN

Accession

Primary (citable) accession number: Q96P70
Secondary accession number(s): B1ASV5 , Q8N1Y1, Q8N3I2, Q8NCG9, Q96SU6, Q9NW01, Q9P0A8, Q9ULM8

Entry history

Integrated into UniProtKB/Swiss-Prot:	February 2, 2004
Last sequence update:	January 23, 2007
Last modified:	May 1, 2013
This is version 105 of the entry and version 3 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.