Importin

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Structure of yeast importin α NLS-binding domain (grey) complex with nucleoplasmin peptide (green) (PDB code 1ee5).

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References

↑ Mattaj IW, Englmeier L. Nucleocytoplasmic transport: the soluble phase. Annu Rev Biochem. 1998;67:265-306. PMID:9759490 doi:http://dx.doi.org/10.1146/annurev.biochem.67.1.265
↑ Bernardes NE, Fung HYJ, Li Y, Chen Z, Chook YM. Structure of IMPORTIN-4 bound to the H3-H4-ASF1 histone-histone chaperone complex. Proc Natl Acad Sci U S A. 2022 Sep 20;119(38):e2207177119. PMID:36103578 doi:10.1073/pnas.2207177119
↑ García-García M, Sánchez-Perales S, Jarabo P, Calvo E, Huyton T, Fu L, Ng SC, Sotodosos-Alonso L, Vázquez J, Casas-Tintó S, Görlich D, Echarri A, Del Pozo MA. Mechanical control of nuclear import by Importin-7 is regulated by its dominant cargo YAP. Nat Commun. 2022 Mar 4;13(1):1174. PMID:35246520 doi:10.1038/s41467-022-28693-y
↑ Padavannil A, Sarkar P, Kim SJ, Cagatay T, Jiou J, Brautigam CA, Tomchick DR, Sali A, D'Arcy S, Chook YM. Importin-9 wraps around the H2A-H2B core to act as nuclear importer and histone chaperone. Elife. 2019 Mar 11;8. pii: 43630. doi: 10.7554/eLife.43630. PMID:30855230 doi:http://dx.doi.org/10.7554/eLife.43630
↑ Gajewska KA, Lescesen H, Ramialison M, Wagstaff KM, Jans DA. Nuclear transporter Importin-13 plays a key role in the oxidative stress transcriptional response. Nat Commun. 2021 Oct 8;12(1):5904. PMID:34625540 doi:10.1038/s41467-021-26125-x
↑ Pumroy RA, Cingolani G. Diversification of importin-alpha isoforms in cellular trafficking and disease states. Biochem J. 2015 Feb 15;466(1):13-28. doi: 10.1042/BJ20141186. PMID:25656054 doi:http://dx.doi.org/10.1042/BJ20141186

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Importin

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@@ Line 3: / Line 3: @@
 == Function ==
-'''Importins''' (IPO) mediate the import of cellular proteins into the nucleus by binding to the '''N'''uclear '''L'''ocalization '''S'''ignal (NLS).  IPO has 2 subunits: α and β.  '''IPO-α''' is an adaptor protein which binds the NLS<ref>PMID:9759490</ref>.  IPOs can contain one or two NLS motifs.  '''IPO-α''' contains several armadillo repeats with two NLS-binding sites and an IPO-β binding (IBB) site.  '''IPO-β''' contains 19 HEAT (solenoid domain) repeats.  Many nuclear proteins contain Phe-Gly sequences which can bind to HEAT repeats and thus be transported.  See also [[Protein Transport Membrane Protein]].
+'''Importins''' (IPO) mediate the import of cellular proteins into the nucleus by binding to the '''N'''uclear '''L'''ocalization '''S'''ignal (NLS).  IPO has 2 subunits: α and β.  '''IPO-α''' is an adaptor protein which binds the NLS<ref>PMID:9759490</ref>.  IPOs can contain one or two NLS motifs.
+*'''IPO-α''' contains several armadillo repeats with two NLS-binding sites and an IPO-β binding (IBB) site.
+*'''IPO-β''' contains 19 HEAT (solenoid domain) repeats.  Many nuclear proteins contain Phe-Gly sequences which can bind to HEAT repeats and thus be transported.
+*'''IPO-4''' binds histones H3 and H4 and histone chaperone ASF1 for nuclear import<ref>PMID:36103578</ref>.
+*'''IPO-7'''  drives the nuclear import of the regulator of mechanotransduction pathway YAP<ref>PMID:35246520</ref>.
+*'''IPO-9''' binds histones H2A and H2B for nuclear import<ref>PMID:30855230</ref>.
+*'''IPO-13''' has a role in oxidative stress-dependent nuclear export<ref>PMID:34625540</ref>.
+See also [[Protein Transport Membrane Protein]].
 == Disease ==
@@ Line 17: / Line 25: @@
 </StructureSection>
-==3D structures of importin==
-Updated on {{REVISIONDAY2}}-{{MONTHNAME|{{REVISIONMONTH}}}}-{{REVISIONYEAR}}
-{{#tree:id=OrganizedByTopic|openlevels=0|
-*IPO-α
-**[[1ial]] – mIPO NLS-binding domain – mouse<br />
-**[[1bk5]] – yIPO NLS-binding domain - yeast<br />
-**[[4tnm]] – IPO armadillo repeat domain – ''Arabidopsis thaliana''<br />
-*''IPO-α complex with NLS peptide''
-**[[1ejl]] – mIPO NLS-binding domain + SV40 large T antigen<br />
-**[[1ejy]] – mIPO NLS-binding domain + nucleoplasmin <br />
-**[[2ynr]] – mIPO NLS-binding domain + B54NLS <br />
-**[[2yns]] – rIPO NLS-binding domain + B54NLS - rice<br />
-**[[1bk6]] – yIPO NLS-binding domain + SV40 large T antigen<br />
-**[[1ee4]] – yIPO NLS-binding domain (mutant) + c-Myc<br />
-**[[1ee5]] – yIPO NLS-binding domain (mutant) + nucleoplasmin<br />
-**[[1un0]], [[2c1t]] – yIPO NLS-binding domain (mutant) + Nup2<br />
-**[[5h2x]], [[5h2w]] – yIPO NLS-binding domain + ubiquitin-like specific protease<br />
-**[[4xzr]] – yIPO NLS-binding domain + HEH1<br />
-**[[4pvz]] – yIPO NLS-binding domain + HEH2<br />
-**[[4rxh]] – NcIPO NLS-binding domain + SV40 large T antigen – ''Neurospora crassa''<br />
-**[[5vqi]] – NcIPO NLS-binding domain + NIT2<br />
-*''IPO-α complex with protein''
-**[[1wa5]] – yIPO NLS-binding domain + GTPase Ran + IPO-α re-exporter<br />
-**[[5t94]] – yIPO NLS-binding domain + guanine nucleotide exchange factor <br />
-**[[5mfm]] – yIPO NLS-binding domain + YIIIM6AII <br />
-*IPO-α 1
-**[[3wpt]] – hIPO NLS-binding domain - human<br />
-**[[4b8j]] – rIPO <br />
-**[[6bvt]] – mIPO NLS-binding domain <br />
-*''IPO-α 1 complex with NLS peptide''
-**[[4b18]] – hIPO NLS-binding domain + telomerase reverse transcriptase<br />
-**[[4wv6]] – hIPO NLS-binding domain + TFIID<br />
-**[[6bw0]] – hIPO NLS-binding domain + Nipa virus peptide<br />
-**[[6bw1]] – hIPO NLS-binding domain + Hendra virus peptide<br />
-**[[5w4e]], [[5w4f]], [[5w4g]] – hIPO NLS-binding domain + polymerase peptide<br />
-**[[5h43]] – hIPO NLS-binding domain + histone acetyltransferase KAT8<br />
-**[[4mz5]], [[4mz6]] – mIPO NLS-binding domain + dUTPase<br />
-**[[4oih]] – mIPO NLS-binding domain + guanine nucleotide exchange factor SRM1<br />
-**[[5svz]] – mIPO NLS-binding domain + TAT NLS motif<br />
-**[[5wum]], [[5wun]], [[5x8n]] – mIPO NLS-binding domain + Epstein Barr antigen<br />
-**[[5huw]] – mIPO NLS-binding domain + HSVI large terminase<br />
-**[[5huy]] – mIPO NLS-binding domain + HSMV small terminase<br />
-**[[5hhg]], [[5fc8]] – mIPO NLS-binding domain + dengue 2 NS5<br />
-**[[5ekg]], [[5ekf]], [[5e6q]] – mIPO NLS-binding domain + XPG2<br />
-**[[5d5k]] – mIPO NLS-binding domain + PARP 2<br />
-**[[5ctt]] – mIPO NLS-binding domain + SART3/TIP110<br />
-**[[5b56]] – mIPO NLS-binding domain + HIV-1 VPR<br />
-**[[4zdu]] – mIPO NLS-binding domain + influenza nucleoprotein<br />
-**[[4yi0]] – mIPO NLS-binding domain + Nup121<br />
-**[[5klr]], [[5klt]] – mIPO NLS-binding domain + NLS <br />
-**[[5k9s]] – mIPO NLS-binding domain + HNF-b <br />
-**[[6iu7]], [[6iua]] – mIPO NLS-binding domain + TP53-binding protein <br />
-**[[5w41]] – mIPO NLS-binding domain + Zika virus peptide <br />
-**[[5v5o]], [[5v5p]] – mIPO NLS-binding domain + influenza virus peptide <br />
-**[[5umz]] – mIPO NLS-binding domain + TNRC6A peptide <br />
-**[[5u5p]], [[5u5r]] – mIPO NLS-binding domain + MLH1 peptide <br />
-**[[5gxw]] – mIPO NLS-binding domain + NUMA peptide <br />
-**[[4b8o]] – rIPO NLS-binding domain + SV40<br />
-**[[4b8p]] – rIPO NLS-binding domain + A89<br />
-**[[4bpl]] – rIPO NLS-binding domain + nucleoplasmin<br />
-**[[4bqk]] – rIPO NLS-binding domain + VirD2<br />
-*''IPO-α 1 complex with protein''
-**[[2jdq]] – hIPO NLS-binding domain + polymerase basic protein  2 C terminal<br />
-**[[3tj3]] – hIPO NLS-binding domain + Nup50 N terminal<br />
-**[[2jdq]] – hIPO NLS-binding domain + telomerase reverse transcriptase peptide<br />
-**[[4uaf]] – mIPO NLS-binding domain + influenza RNA polymerase<br />
-*IPO-α 2
-**[[3tpo]] – mIPO<br />
-**[[1iq1]] – mIPO NLS-binding + autoinhibitory peptide domains<br />
-**[[4e4v]] – hIPO NLS-binding domain (mutant) <br />
-*''IPO-α 2 complex with NLS peptide''
-**[[1pjm]] – mIPO NLS-binding domain + retinoblastoma-associated protein<br />
-**[[1pjn]] – mIPO NLS-binding domain + histone-binding protein N1/N2<br />
-**[[1q1s]], [[1q1t]] – mIPO NLS-binding domain + SV40 large T antigen<br />
-**[[1y2a]], [[3q5u]] – mIPO NLS-binding domain + phospholipid scramblase <br />
-**[[2c1m]] – mIPO NLS-binding domain + Nup50<br />
-**[[3btr]] – mIPO NLS-binding domain + androgen receptor<br />
-**[[3knd]] – mIPO NLS-binding domain + targeting protein for KXLP2<br />
-**[[3l3q]] – mIPO NLS-binding domain + library peptide<br />
-**[[3oqs]] – mIPO NLS-binding domain + CLIC4<br />
-**[[3rz9]] – mIPO NLS-binding domain + KU80<br />
-**[[3rzx]] – mIPO NLS-binding domain + KU70<br />
-**[[3tpm]] – mIPO NLS-binding domain + MAL<br />
-**[[3ve6]] – mIPO NLS-binding domain + Venezuelan equine encephalitis virus<br />
-**[[3uvu]] – mIPO NLS-binding domain + flap endonuclease<br />
-**[[3ukw]], [[3ukx]] – mIPO NLS-binding domain + BIMAX<br />
-**[[3uky]], [[3ukz]], [[3ul0]] – mIPO NLS-binding domain + nuclear CAP-binding protein<br />
-**[[3ul1]] – mIPO NLS-binding domain + nucleoplasmin <br />
-**[[4htv]] – mIPO NLS-binding domain + capsid protein <br />
-**[[4ba3]] – mIPO NLS-binding domain + A89NLS <br />
-**[[3zio]], [[3zip]], [[3ziq]], [[3zir]] – mIPO NLS-binding domain + minor site-specific NLS <br />
-**[[3zin]] – mIPO NLS-binding domain + nucleolar RNA helicase<br />
-*''IPO-α 2 complex with protein''
-**[[3fex]], [[3fey]] – mIPO NLS-binding domain + nuclear CAP-binding protein<br />
-*IPO-α 3
-**[[6bvz]] – hIPO NLS-binding domain <br />
-**[[4uae]] – hIPO NLS-binding domain + influenza RNA polymerase<br />
-**[[6bw9]], [[6bwa]], [[6bwb]] – hIPO NLS-binding domain + Hendra virus peptide<br />
-**[[6bvv]] – hIPO NLS-binding domain + Nipa virus peptide<br />
-**[[5xzx]] – hIPO NLS-binding domain + RAN-binding protein peptide<br />
-**[[5tbk]] – hIPO NLS-binding domain + regulator of chromosome condensation <br />
-*IPO-α 6
-**[[4u2x]] – hIPO C terminal + ebola VP24<br />
-*IPO-α 7
-**[[4uad]] – hIPO NLS-binding domain + influenza RNA polymerase<br />
-*IPO-β
-**[[4xrk]] – IPO – ''Chaetomium thermophilum'' <br />
-*IPO-β 1 (karyopherin β 1)
-**[[3nd2]] – yIPO <br />
-**[[1gcj]] – mIPO N terminal<br />
-*''IPO-β 1 complex with protein''
-**[[1qgx]], [[1qgk]] – hIPO + IPO-α 2 IBB domain<br />
-**[[1ibr]] – hIPO + GTPase Ran<br />
-**[[2bku]], [[3ea5]] – yIPO + GTPase Ran<br />
-**[[1f59]], [[1o6p]], [[1o6o]] – hIPO + FXFG nucleoporin repeats<br />
-**[[1m5n]] – hIPO HEAT repeats 1-11 + parathyroid hormone-related protein NLS peptide<br />
-**[[2p8q]], [[2q5d]], [[2qna]], [[3lww]] – hIPO + RNA U transporter N terminal<br />
-**[[1qgr]], [[3w5k]] – hIPO + zinc finger protein SNAI1<br />
-**[[5owu]] – yIPO + Nup1<br />
-**[[1ukl]] – mIPO + SREBP-2<br />
-*IPO-β 2 (Transportin 1)
-**[[2z5j]], [[2qmr]] – hIPO <br />
-*''IPO-β 2 complex with protein''
-**[[2h4m]], [[2ot8]], [[2z5n]] – hIPO + hnRNP peptide<br />
-**[[2z5k]], [[2z5m]] – hIPO + nuclear RNA export factor peptide<br />
-**[[2z5o]] – hIPO + JKTBP peptide<br />
-**[[4fdd]], [[4jlq]], [[4oo6]] – hIPO + RNA binding protein peptide<br />
-**[[5yvg]], [[5yvh]] – hIPO + FUS <br />
-**[[5vyh]], [[5vyi]] – hIPO + FUS peptide<br />
-**[[4fq3]] – hIPO + FUS peptide<br />
-**[[5tqc]] – hIPO + RPL4 peptide<br />
-**[[5j3v]] – hIPO + histone H3 peptide<br />
-*IPO-β 3 (Karyopherin β 3)
-**[[3w3t]] – yIPO <br />
-**[[3w3u]], [[3w3v]] – yIPO (mutant)<br />
-**[[3w3w]] – yIPO + STE12P<br />
-**[[3w3x]] – yIPO + PHO4<br />
-**[[3w3y]] – yIPO + Nup53P<br />
-**[[3w3z]] – yIPO + RANGTP<br />
-**[[5h2v]] – yIPO + ubiquitin-like specific protease<br />
-**[[4zj7]] – yIPO + CDC14<br />
-*IPO-4B
-**[[5xah]] – hIPO residues 668-1081  <br />
-**[[5xbk]] – hIPO residues 668-1081 + histone H3 peptide <br />
-*IPO-7
-**[[6n88]] – IPO + hIPO-90 + histone H1.0 – frog – Cryo EM  <br />
-*IPO-9
-**[[6n1z]] – hIPO + histone H2A + histone H2B  <br />
-*IPO-12 (Transportin-3)
-**[[4c0p]] – hIPO <br />
-**[[4c0o]] – hIPO + serine/arginine-rich splicing factor<br />
-**[[4c0q]], [[4ol0]] – hIPO + GTPase Ran<br />
-*IPO-13 (karyopherin 13)
-**[[2x19]] – hIPO + GTPase Ran<br />
-**[[2xwu]] – hIPO + UBC9<br />
-**[[3zjy]] – hIPO + GTPase Ran + EIF-1A<br />
-*IPO-90
-**[[6n89]] – hIPO-90 + histone H1.0 – Cryo EM  <br />
-}}
 == References ==