beta karyopherins


Summary: Nucleocytoplasmic transport molecules that bind to ALPHA KARYOPHERINS in the CYTOSOL and are involved in transport of molecules through the NUCLEAR PORE COMPLEX. Once inside the CELL NUCLEUS beta karyopherins interact with RAN GTP-BINDING PROTEIN and dissociate from alpha karyopherins. Beta karyopherins bound to RAN GTP-BINDING PROTEIN are then re-transported to the cytoplasm where hydrolysis of the GTP of RAN GTP-BINDING PROTEIN causes release of karyopherin beta.

Top Publications

  1. Ryan K, Zhou Y, Wente S. The karyopherin Kap95 regulates nuclear pore complex assembly into intact nuclear envelopes in vivo. Mol Biol Cell. 2007;18:886-98 pubmed
    ..We conclude that Kap95 serves as a key regulator of NPC assembly into intact NEs. Furthermore, both Kap95 and Ran may provide spatial cues necessary for targeting of vesicular intermediates in de novo NPC assembly. ..
  2. Dias S, Wilson K, Rojas K, Ambrosio A, Cerione R. The molecular basis for the regulation of the cap-binding complex by the importins. Nat Struct Mol Biol. 2009;16:930-7 pubmed publisher
    ..Together, these studies enable us to propose a model describing how importin-beta stimulates the dissociation of capped RNA from the CBC in the cytosol following its nuclear export. ..
  3. Pelaez R, Fernández García P, Herrero P, Moreno F. Nuclear import of the yeast hexokinase 2 protein requires ?/?-importin-dependent pathway. J Biol Chem. 2012;287:3518-29 pubmed publisher
    ..It is also demonstrated that Hxk2 nuclear import and its binding to Kap95 and Kap60 depend on the Gsp1-GTP/GDP protein levels. Thus, our study uncovers Hxk2 as a new cargo for the ?/?-importin pathway of S. cerevisiae. ..
  4. Yang W, Musser S. Nuclear import time and transport efficiency depend on importin beta concentration. J Cell Biol. 2006;174:951-61 pubmed
  5. Logue E, Taylor K, Goff P, Landau N. The cargo-binding domain of transportin 3 is required for lentivirus nuclear import. J Virol. 2011;85:12950-61 pubmed publisher
    ..The mutated TNPO3 proteins maintained their ability to localize to the nucleus, suggesting that their inability to restore lentivirus infection resulted from an inability to bind to a host or viral cargo protein. ..
  6. Levin A, Hayouka Z, Friedler A, Loyter A. Transportin 3 and importin ? are required for effective nuclear import of HIV-1 integrase in virus-infected cells. Nucleus. 2010;1:422-31 pubmed publisher
  7. Süel K, Chook Y. Kap104p imports the PY-NLS-containing transcription factor Tfg2p into the nucleus. J Biol Chem. 2009;284:15416-24 pubmed publisher
    ..More generally, steady-state localization of a nuclear protein is dictated by its nuclear import and export activities as well as its interactions in the nucleus and the cytoplasm. ..
  8. Rebane A, Aab A, Steitz J. Transportins 1 and 2 are redundant nuclear import factors for hnRNP A1 and HuR. RNA. 2004;10:590-9 pubmed
    ..Possible nucleocytoplasmic shuttling mechanisms for hnRNP A1 and HuR are discussed. ..
  9. Ocwieja K, Brady T, Ronen K, Huegel A, Roth S, Schaller T, et al. HIV integration targeting: a pathway involving Transportin-3 and the nuclear pore protein RanBP2. PLoS Pathog. 2011;7:e1001313 pubmed publisher
    ..Thus, our data support a model in which Gag-dependent engagement of the proper transport and nuclear pore machinery mediate trafficking of HIV complexes to sites of integration. ..

More Information


  1. Cushman I, Palzkill T, Moore M. Using peptide arrays to define nuclear carrier binding sites on nucleoporins. Methods. 2006;39:329-41 pubmed
  2. Quensel C, Friedrich B, Sommer T, Hartmann E, Kohler M. In vivo analysis of importin alpha proteins reveals cellular proliferation inhibition and substrate specificity. Mol Cell Biol. 2004;24:10246-55 pubmed
    ..Thus, importin alpha3 is the only alpha importin responsible for the classical nuclear import of RCC1 in living cells. ..
  3. Dishinger J, Kee H, Jenkins P, Fan S, Hurd T, Hammond J, et al. Ciliary entry of the kinesin-2 motor KIF17 is regulated by importin-beta2 and RanGTP. Nat Cell Biol. 2010;12:703-10 pubmed publisher
    ..We propose that Ran has a global role in regulating cellular compartmentalization by controlling the shuttling of cytoplasmic proteins into nuclear and ciliary compartments. ..
  4. Yasumi M, Sakisaka T, Hoshino T, Kimura T, Sakamoto Y, Yamanaka T, et al. Direct binding of Lgl2 to LGN during mitosis and its requirement for normal cell division. J Biol Chem. 2005;280:6761-5 pubmed
    ..NuMA complex. These results indicate that Lgl2 forms a Lgl2.Par-6.aPKC.LGN complex, which responds to mitotic signaling to establish normal cell division. ..
  5. Lim R, Fahrenkrog B, Köser J, Schwarz Herion K, Deng J, Aebi U. Nanomechanical basis of selective gating by the nuclear pore complex. Science. 2007;318:640-3 pubmed
    ..Similar effects were observed in Xenopus oocyte nuclei in situ. Thus, the reversible collapse of the FG domains may play an important role in regulating nucleocytoplasmic transport. ..
  6. Kubitscheck U, GRUNWALD D, Hoekstra A, Rohleder D, Kues T, Siebrasse J, et al. Nuclear transport of single molecules: dwell times at the nuclear pore complex. J Cell Biol. 2005;168:233-43 pubmed
    ..Together with the known transport rates, our data suggest that nucleocytoplasmic transport occurs via multiple parallel pathways within single NPCs. ..
  7. Frankel M, Knoll L. The ins and outs of nuclear trafficking: unusual aspects in apicomplexan parasites. DNA Cell Biol. 2009;28:277-84 pubmed publisher
    ..A detailed analysis of the nuclear trafficking network in these eukaryotes is required to understand how this potentially unique cellular biological pathway contributes to host-parasite interactions. ..
  8. Lange A, Mills R, Devine S, Corbett A. A PY-NLS nuclear targeting signal is required for nuclear localization and function of the Saccharomyces cerevisiae mRNA-binding protein Hrp1. J Biol Chem. 2008;283:12926-34 pubmed publisher
    ..Finally, we apply this analysis to a bioinformatic search of the yeast proteome as a preliminary search for new potential Kap104 cargos. ..
  9. Zhang H, Ju G, Wei J, Hu Y, Liu L, Xu Q, et al. A combined effect of the KPNA3 and KPNB3 genes on susceptibility to schizophrenia. Neurosci Lett. 2006;402:173-5 pubmed
    ..79, P=0.001) and for the rs626716(C)-rs3736830(G) combination (X2=8.64, P=0.003). The present work suggests that the combination of the KPNA3 gene and the KPNB3 gene may increase a genetic risk for schizophrenia. ..
  10. Cardarelli F, Bizzarri R, Serresi M, Albertazzi L, Beltram F. Probing nuclear localization signal-importin alpha binding equilibria in living cells. J Biol Chem. 2009;284:36638-46 pubmed publisher
    ..Finally, our experiments also provide evidence for the regulation of nuclear import mediated by the intrasteric importin beta-binding domain of Impalpha and yield the first estimate of its autoinhibition energy in living cells. ..
  11. Chou C, Tai L, Kirby R, Lee I, Lin A. Importin ?3 mediates the nuclear import of human ribosomal protein L7 through its interaction with the multifaceted basic clusters of L7. FEBS Lett. 2010;584:4151-6 pubmed publisher
    ..We assume that having such a multiple NLS may provide L7 with preferential nuclear uptake. ..
  12. Ghildyal R, Ho A, Wagstaff K, Dias M, Barton C, Jans P, et al. Nuclear import of the respiratory syncytial virus matrix protein is mediated by importin beta1 independent of importin alpha. Biochemistry. 2005;44:12887-95 pubmed
  13. Mehta T, Lu H, Wang X, Urvalek A, Nguyen K, Monzur F, et al. A unique sequence in the N-terminal regulatory region controls the nuclear localization of KLF8 by cooperating with the C-terminal zinc-fingers. Cell Res. 2009;19:1098-109 pubmed publisher
    ..Taken together, these results suggest that KLF8 has two NLSs, one surrounding S165 and K171 and the other being two tandem ZFs, which are critical for the regulation of KLF8 nuclear localization and its cellular functions. ..
  14. Zhou L, Sokolskaja E, Jolly C, James W, Cowley S, Fassati A. Transportin 3 promotes a nuclear maturation step required for efficient HIV-1 integration. PLoS Pathog. 2011;7:e1002194 pubmed publisher
    ..The results also provide evidence for a novel tRNA nucleocytoplasmic trafficking pathway in human cells. ..
  15. Tsai M, Wang S, Heidinger J, Shumaker D, Adam S, Goldman R, et al. A mitotic lamin B matrix induced by RanGTP required for spindle assembly. Science. 2006;311:1887-93 pubmed
    ..We propose that lamin B is a structural component of the long-sought-after spindle matrix that promotes microtubule assembly and organization in mitosis. ..
  16. Taberner F, Igual J. Yeast karyopherin Kap95 is required for cell cycle progression at Start. BMC Cell Biol. 2010;11:47 pubmed publisher
    ..This transport depends on the specific nuclear localization signals present in cargo proteins. ..
  17. Yamasaki H, Sekimoto T, Ohkubo T, Douchi T, Nagata Y, Ozawa M, et al. Zinc finger domain of Snail functions as a nuclear localization signal for importin beta-mediated nuclear import pathway. Genes Cells. 2005;10:455-64 pubmed
    ..These results indicate that zinc finger domain of Snail functions as a nuclear localization signal and Snail can be transported into the nucleus in an importin beta-mediated manner. ..
  18. Isgro T, Schulten K. Binding dynamics of isolated nucleoporin repeat regions to importin-beta. Structure. 2005;13:1869-79 pubmed
  19. Ems McClung S, Zheng Y, Walczak C. Importin alpha/beta and Ran-GTP regulate XCTK2 microtubule binding through a bipartite nuclear localization signal. Mol Biol Cell. 2004;15:46-57 pubmed
    ..Our data show that importin alpha/beta directly regulates the activity of XCTK2 and that one of the molecular mechanisms of Ran-regulated spindle assembly is identical to that used in classical NLS-driven nuclear transport. ..
  20. Chook Y, Süel K. Nuclear import by karyopherin-?s: recognition and inhibition. Biochim Biophys Acta. 2011;1813:1593-606 pubmed publisher
    ..This article is part of a Special Issue entitled: Regulation of Signaling and Cellular Fate through Modulation of Nuclear Protein Import. ..
  21. Süel K, Gu H, Chook Y. Modular organization and combinatorial energetics of proline-tyrosine nuclear localization signals. PLoS Biol. 2008;6:e137 pubmed publisher
    ..The modular organization of the PY-NLS coupled with its combinatorial energetics lays a path to decode this diverse and evolvable signal for future comprehensive genome-scale identification of nuclear import substrates. ..
  22. Shah V, Shi J, Hout D, Oztop I, Krishnan L, Ahn J, et al. The host proteins transportin SR2/TNPO3 and cyclophilin A exert opposing effects on HIV-1 uncoating. J Virol. 2013;87:422-32 pubmed publisher
    ..Our results suggest that TNPO3 and cyclophilin A facilitate HIV-1 infection by coordinating proper uncoating of the core in target cells. ..
  23. Kumari G, Singhal P, Rao M, Mahalingam S. Nuclear transport of Ras-associated tumor suppressor proteins: different transport receptor binding specificities for arginine-rich nuclear targeting signals. J Mol Biol. 2007;367:1294-311 pubmed
    ..Together, these data suggest that the transport of Ras effector superfamily proteins into the nucleus/nucleolus may play a vital role in modulating Ras-mediated cell proliferation during tumorigenesis. ..
  24. Kalab P, Pralle A, Isacoff E, Heald R, Weis K. Analysis of a RanGTP-regulated gradient in mitotic somatic cells. Nature. 2006;440:697-701 pubmed
  25. Sakakida Y, Miyamoto Y, Nagoshi E, Akashi M, Nakamura T, Mamine T, et al. Importin alpha/beta mediates nuclear transport of a mammalian circadian clock component, mCRY2, together with mPER2, through a bipartite nuclear localization signal. J Biol Chem. 2005;280:13272-8 pubmed
    ..These results suggest that the importin alpha/beta system is involved in nuclear entry of mammalian clock components, which is indispensable to transcriptional oscillation of clock genes. ..
  26. Petosa C, Schoehn G, Askjaer P, Bauer U, Moulin M, Steuerwald U, et al. Architecture of CRM1/Exportin1 suggests how cooperativity is achieved during formation of a nuclear export complex. Mol Cell. 2004;16:761-75 pubmed
    ..Our findings indicate that a single architectural and mechanistic framework can explain the divergent effects of RanGTP on substrate binding by many import and export receptors. ..
  27. Schmitz M, Held M, Janssens V, Hutchins J, Hudecz O, Ivanova E, et al. Live-cell imaging RNAi screen identifies PP2A-B55alpha and importin-beta1 as key mitotic exit regulators in human cells. Nat Cell Biol. 2010;12:886-93 pubmed publisher
    ..This demonstrates that PP2A-B55alpha and importin-beta1 cooperate in the regulation of postmitotic assembly mechanisms in human cells. ..
  28. Leslie D, Zhang W, Timney B, Chait B, Rout M, Wozniak R, et al. Characterization of karyopherin cargoes reveals unique mechanisms of Kap121p-mediated nuclear import. Mol Cell Biol. 2004;24:8487-503 pubmed
    ..Together, our data elucidate additional levels of complexity in these nuclear transport pathways. ..
  29. Ploski J, Shamsher M, Radu A. Paired-type homeodomain transcription factors are imported into the nucleus by karyopherin 13. Mol Cell Biol. 2004;24:4824-34 pubmed
    ..All members contain a region similar to the NLS found in Pax6 and are therefore likely to be imported by Kap13. We confirmed this hypothesis for Pax3 and Crx, which bind to and are imported by Kap13. ..
  30. Rotem A, Gruber R, Shorer H, Shaulov L, Klein E, Harel A. Importin beta regulates the seeding of chromatin with initiation sites for nuclear pore assembly. Mol Biol Cell. 2009;20:4031-42 pubmed publisher
    ..We predict that additional regulators, besides importin beta and Ran, may be involved in coordinating the initial seeding of chromatin with subsequent steps in the NPC assembly pathway. ..
  31. Van Dusen C, Yee L, McNally L, McNally M. A glycine-rich domain of hnRNP H/F promotes nucleocytoplasmic shuttling and nuclear import through an interaction with transportin 1. Mol Cell Biol. 2010;30:2552-62 pubmed publisher
    ..Collectively, the results suggest that hnRNP H and F are GYR domain-dependent shuttling proteins whose posttranslational modifications may alter nuclear localization and hence function. ..
  32. Song L, Rape M. Regulated degradation of spindle assembly factors by the anaphase-promoting complex. Mol Cell. 2010;38:369-82 pubmed publisher
    ..Our findings reveal a tightly regulated mechanism by which the APC/C and the GTPase Ran control the abundance of active spindle assembly factors to achieve the accurate formation of the mitotic spindle. ..
  33. Depping R, Steinhoff A, Schindler S, Friedrich B, Fagerlund R, Metzen E, et al. Nuclear translocation of hypoxia-inducible factors (HIFs): involvement of the classical importin alpha/beta pathway. Biochim Biophys Acta. 2008;1783:394-404 pubmed publisher
    ..In contrast, the supposed N-terminal NLS is not effective. Our findings provide new insight into the mechanism of the regulation of nuclear transport of hypoxia-inducible factors. ..
  34. Hamada M, Haeger A, Jeganathan K, van Ree J, Malureanu L, Wälde S, et al. Ran-dependent docking of importin-beta to RanBP2/Nup358 filaments is essential for protein import and cell viability. J Cell Biol. 2011;194:597-612 pubmed publisher
    ..These data suggest that a critical function of RanBP2 is to capture recycling RanGTP-importin-? complexes at cytoplasmic fibrils to allow for adequate cNLS-mediated cargo import. ..
  35. Fan S, Fogg V, Wang Q, Chen X, Liu C, Margolis B. A novel Crumbs3 isoform regulates cell division and ciliogenesis via importin beta interactions. J Cell Biol. 2007;178:387-98 pubmed
    ..Knockdown of importin beta-1 blocks targeting of CRB3-CLPI to the spindle poles. Our data suggest an expanded role for Crumbs proteins in polarized membrane targeting and cell division via unique interactions with importin proteins. ..
  36. Larue R, Gupta K, Wuensch C, Shkriabai N, Kessl J, Danhart E, et al. Interaction of the HIV-1 intasome with transportin 3 protein (TNPO3 or TRN-SR2). J Biol Chem. 2012;287:34044-58 pubmed publisher
    ..Our findings provide important information for future genetic analysis to better understand the role of TNPO3 and its interacting partners for HIV-1 replication. ..
  37. Lange A, Mills R, Lange C, Stewart M, Devine S, Corbett A. Classical nuclear localization signals: definition, function, and interaction with importin alpha. J Biol Chem. 2007;282:5101-5 pubmed
    ..Finally, we describe how a predicted NLS within a protein of interest can be confirmed experimentally to be functionally important. ..
  38. Albee A, Tao W, Wiese C. Phosphorylation of maskin by Aurora-A is regulated by RanGTP and importin beta. J Biol Chem. 2006;281:38293-301 pubmed
    ..We further show that importin beta inhibits the regulatory phosphorylation of maskin by Aurora-A. This suggests a novel mechanism by which importin beta regulates the activity of a spindle assembly factor. ..
  39. Merkle T. Nucleo-cytoplasmic partitioning of proteins in plants: implications for the regulation of environmental and developmental signalling. Curr Genet. 2003;44:231-60 pubmed
    ..Finally, the importance of nucleo-cytoplasmic partitioning as a regulatory tool for signalling is highlighted, and different plant signal transduction pathways that make use of this regulatory potential are presented. ..
  40. Koike M, Kose S, Furuta M, Taniguchi N, Yokoya F, Yoneda Y, et al. beta-Catenin shows an overlapping sequence requirement but distinct molecular interactions for its bidirectional passage through nuclear pores. J Biol Chem. 2004;279:34038-47 pubmed
    ..The data herein indicate that beta-catenin shows an overlapping sequence requirement for its import and export but that bidirectional movement through the NPC proceeds through distinct molecular interactions. ..
  41. Yasuhara N, Takeda E, Inoue H, Kotera I, Yoneda Y. Importin alpha/beta-mediated nuclear protein import is regulated in a cell cycle-dependent manner. Exp Cell Res. 2004;297:285-93 pubmed
    ..These results indicate that the importin alpha/beta-mediated nuclear import machinery is regulated in a cell cycle-dependent manner through the modulation of interaction modes between importins alpha and beta. ..
  42. Zhang Y, Sperry A. Comparative analysis of two C-terminal kinesin motor proteins: KIFC1 and KIFC5A. Cell Motil Cytoskeleton. 2004;58:213-30 pubmed
    ..The KIFC5A tail contains a 43 amino acid sequence with both nuclear localization and microtubule binding activity while KIFC1 contains a 19 amino acid sequence sufficient to target this motor to membrane-bounded organelles. ..
  43. Goldfarb D, Corbett A, Mason D, Harreman M, Adam S. Importin alpha: a multipurpose nuclear-transport receptor. Trends Cell Biol. 2004;14:505-14 pubmed
    ..We also describe activities of importin alpha that seem to be distinct from its housekeeping functions in nuclear transport. ..
  44. Harel A, Forbes D. Importin beta: conducting a much larger cellular symphony. Mol Cell. 2004;16:319-30 pubmed
    ..Lastly, importin beta plays a role in transducing damage signals from the axons of injured neurons back to the cell body. ..
  45. Plafker S, Plafker K, Weissman A, Macara I. Ubiquitin charging of human class III ubiquitin-conjugating enzymes triggers their nuclear import. J Cell Biol. 2004;167:649-59 pubmed
    ..Together, these findings reveal that Ub charging can function as a nuclear import trigger, and identify a novel link between E2 regulation and karyopherin-mediated transport. ..
  46. Blower M, Nachury M, Heald R, Weis K. A Rae1-containing ribonucleoprotein complex is required for mitotic spindle assembly. Cell. 2005;121:223-34 pubmed
    ..Furthermore, we provide evidence that RNA associates with the mitotic spindle and that it plays a direct, translation-independent role in spindle assembly. Our studies reveal an unexpected function for RNA in spindle morphogenesis. ..
  47. Lee S, Matsuura Y, Liu S, Stewart M. Structural basis for nuclear import complex dissociation by RanGTP. Nature. 2005;435:693-6 pubmed
    ..This interaction produces a change in helicoidal pitch that locks Kap95p in a conformation that cannot bind importin-alpha or cargo. We suggest an allosteric mechanism for nuclear import complex disassembly by RanGTP. ..
  48. Hearps A, Jans D. HIV-1 integrase is capable of targeting DNA to the nucleus via an importin alpha/beta-dependent mechanism. Biochem J. 2006;398:475-84 pubmed
    ..These results may not only aid in highlighting potential therapeutic targets for impeding the progression of HIV/AIDS, but may also be relevant for non-viral gene delivery. ..
  49. Christ F, Thys W, De Rijck J, Gijsbers R, Albanese A, Arosio D, et al. Transportin-SR2 imports HIV into the nucleus. Curr Biol. 2008;18:1192-202 pubmed publisher
    ..In comparison with control cell lines, the great majority of siRNA-treated cells did not contain any PIC in the nucleus. Our data clearly demonstrate that TRN-SR2 is the nuclear-import factor of HIV. ..
  50. Cribier A, Segeral E, Delelis O, Parissi V, Simon A, Ruff M, et al. Mutations affecting interaction of integrase with TNPO3 do not prevent HIV-1 cDNA nuclear import. Retrovirology. 2011;8:104 pubmed publisher
  51. Quan X, Yu J, Bussey H, Stochaj U. The localization of nuclear exporters of the importin-beta family is regulated by Snf1 kinase, nutrient supply and stress. Biochim Biophys Acta. 2007;1773:1052-61 pubmed
    ..Our results indicate that the association of nuclear exporters with nuclei is controlled by multiple mechanisms that are organized in a hierarchical fashion and linked to the physiological state of the cell. ..
  52. Marfori M, Mynott A, Ellis J, Mehdi A, Saunders N, Curmi P, et al. Molecular basis for specificity of nuclear import and prediction of nuclear localization. Biochim Biophys Acta. 2011;1813:1562-77 pubmed publisher
    ..This article is part of a Special Issue entitled: Regulation of Signaling and Cellular Fate through Modulation of Nuclear Protein Import. ..
  53. Imasaki T, Shimizu T, Hashimoto H, Hidaka Y, Kose S, Imamoto N, et al. Structural basis for substrate recognition and dissociation by human transportin 1. Mol Cell. 2007;28:57-67 pubmed
    ..These studies provide deep understanding of substrate recognition and dissociation by Trn1 in import pathways...
  54. Lee B, Cansizoglu A, Süel K, Louis T, Zhang Z, Chook Y. Rules for nuclear localization sequence recognition by karyopherin beta 2. Cell. 2006;126:543-58 pubmed
    ..These studies define and validate a new NLS that could not be predicted by primary sequence analysis alone. ..
  55. Chalmers A, Pambos M, Mason J, Lang S, Wylie C, Papalopulu N. aPKC, Crumbs3 and Lgl2 control apicobasal polarity in early vertebrate development. Development. 2005;132:977-86 pubmed publisher
    ..We conclude that an instrumental antagonistic interaction between aPKC and Lgl2 defines apicobasal polarity in early vertebrate development...
  56. Tahara K, Takagi M, Ohsugi M, Sone T, Nishiumi F, Maeshima K, et al. Importin-beta and the small guanosine triphosphatase Ran mediate chromosome loading of the human chromokinesin Kid. J Cell Biol. 2008;180:493-506 pubmed publisher
    ..Thus, this study demonstrates a novel nucleocytoplasmic transport factor-mediated mechanism for targeting proteins to mitotic chromosomes. ..
  57. Tao T, Lan J, Lukacs G, Haché R, Kaplan F. Importin 13 regulates nuclear import of the glucocorticoid receptor in airway epithelial cells. Am J Respir Cell Mol Biol. 2006;35:668-80 pubmed
    ..We speculate that variation in cellular levels of IPO13 and intracellular IPO13 shuttling rates may contribute to glucocorticoid resistance. ..
  58. Krishnan L, Matreyek K, Oztop I, Lee K, Tipper C, Li X, et al. The requirement for cellular transportin 3 (TNPO3 or TRN-SR2) during infection maps to human immunodeficiency virus type 1 capsid and not integrase. J Virol. 2010;84:397-406 pubmed publisher
    ..We therefore conclude that capsid, not integrase, is the dominant viral factor that dictates transportin 3 dependency during HIV-1 infection. ..
  59. D Angelo M, Anderson D, Richard E, Hetzer M. Nuclear pores form de novo from both sides of the nuclear envelope. Science. 2006;312:440-3 pubmed
    ..Newly formed nuclear pore complexes did not contain structural components of preexisting pores, suggesting that they can form de novo. ..
  60. Sillje H, Nagel S, Körner R, Nigg E. HURP is a Ran-importin beta-regulated protein that stabilizes kinetochore microtubules in the vicinity of chromosomes. Curr Biol. 2006;16:731-42 pubmed
  61. Kosugi S, Hasebe M, Entani T, Takayama S, Tomita M, Yanagawa H. Design of peptide inhibitors for the importin alpha/beta nuclear import pathway by activity-based profiling. Chem Biol. 2008;15:940-9 pubmed publisher
    ..These peptide inhibitors provide a useful tool for studying nuclear import events. Moreover, our inhibitor design method should enable the development of potent inhibitors from a peptide seed. ..
  62. van der Watt P, Maske C, Hendricks D, Parker M, Denny L, Govender D, et al. The Karyopherin proteins, Crm1 and Karyopherin beta1, are overexpressed in cervical cancer and are critical for cancer cell survival and proliferation. Int J Cancer. 2009;124:1829-40 pubmed publisher
  63. Timney B, Tetenbaum Novatt J, Agate D, Williams R, Zhang W, Chait B, et al. Simple kinetic relationships and nonspecific competition govern nuclear import rates in vivo. J Cell Biol. 2006;175:579-93 pubmed
    ..It is likely that most of every import round is taken up by Kaps and nuclear localization signals sampling other cytoplasmic proteins as they locate each other in the cytoplasm. ..
  64. Ogawa Y, Miyamoto Y, Oka M, Yoneda Y. The interaction between importin-? and Nup153 promotes importin-?/?-mediated nuclear import. Traffic. 2012;13:934-46 pubmed publisher
    ..This is the first in vitro study showing that in conjunction with Nup153, importin ? contributes to directionally biased exit of cNLS-containing cargo to the nuclear side of NPCs. ..
  65. Valle Casuso J, Di Nunzio F, Yang Y, Reszka N, Lienlaf M, Arhel N, et al. TNPO3 is required for HIV-1 replication after nuclear import but prior to integration and binds the HIV-1 core. J Virol. 2012;86:5931-6 pubmed publisher
    ..Overall, this work suggests that TNPO3 interacts with the incoming HIV-1 core in the cytoplasm to assist a process that is important for HIV-1 infection after nuclear import. ..
  66. Tetenbaum Novatt J, Hough L, Mironska R, McKenney A, Rout M. Nucleocytoplasmic transport: a role for nonspecific competition in karyopherin-nucleoporin interactions. Mol Cell Proteomics. 2012;11:31-46 pubmed publisher
  67. Strawn L, Shen T, Shulga N, Goldfarb D, Wente S. Minimal nuclear pore complexes define FG repeat domains essential for transport. Nat Cell Biol. 2004;6:197-206 pubmed
    ..Significantly, symmetric deletions caused mild reductions in Kap95-Kap60-mediated import rates, but virtually abolished Kap104 import. These results suggest the existence of multiple translocation pathways. ..
  68. Saric M, Zhao X, Körner C, Nowak C, Kuhlmann J, Vetter I. Structural and biochemical characterization of the Importin-beta.Ran.GTP.RanBD1 complex. FEBS Lett. 2007;581:1369-76 pubmed
    ..Ran.GTP.RanBD exists in the final step of the export of Importin-beta from the nucleus and that interaction of the N-terminus of RanBD with Ran plays a crucial role in disassembly of this complex. ..
  69. Zhong Y, Wang Y, Yang H, Ballar P, Lee J, Ye Y, et al. Importin beta interacts with the endoplasmic reticulum-associated degradation machinery and promotes ubiquitination and degradation of mutant alpha1-antitrypsin. J Biol Chem. 2011;286:33921-30 pubmed publisher
    ..Furthermore, we show that importin ? cooperates with Ran GTPase to promote ubiquitination and proteasomal degradation of mutant ?1-antitrypsin. These results establish an unanticipated role for importin ? in ER protein quality control. ..
  70. Rao M, Kumari G, Balasundaram D, Sankaranarayanan R, Mahalingam S. A novel lysine-rich domain and GTP binding motifs regulate the nucleolar retention of human guanine nucleotide binding protein, GNL3L. J Mol Biol. 2006;364:637-54 pubmed
    ..Collectively, these data suggest that GNL3L is composed of distinct modules, each of which plays a specific role in molecular interactions for its nucleolar retention and subsequent function(s) within the nucleolus. ..
  71. Belanger K, Griffith A, Baker H, Hansen J, Kovacs L, Seconi J, et al. The karyopherin Kap95 and the C-termini of Rfa1, Rfa2, and Rfa3 are necessary for efficient nuclear import of functional RPA complex proteins in Saccharomyces cerevisiae. DNA Cell Biol. 2011;30:641-51 pubmed publisher
  72. Deng T, Engelhardt O, Thomas B, Akoulitchev A, Brownlee G, Fodor E. Role of ran binding protein 5 in nuclear import and assembly of the influenza virus RNA polymerase complex. J Virol. 2006;80:11911-9 pubmed
    ..Moreover, siRNA knock-down of RanBP5 resulted in the delayed accumulation of viral RNAs in infected cells, confirming that RanBP5 plays a biological role during the influenza virus life cycle. ..
  73. Wei J, Hemmings G. The KPNB3 locus is associated with schizophrenia. Neurosci Lett. 2004;368:323-6 pubmed
    ..18, d.f. = 2, P = 0.006). Because the KPNB3 finding has been replicated in a Chinese population, it could be hypothesized that the KPNB3 locus may contain a disease-causing variant for schizophrenia. ..
  74. Arnold M, Nath A, Hauber J, Kehlenbach R. Multiple importins function as nuclear transport receptors for the Rev protein of human immunodeficiency virus type 1. J Biol Chem. 2006;281:20883-90 pubmed
    ..Taken together, the viral protein takes advantage of multiple cellular transport pathways for its nuclear accumulation. ..
  75. Hu Y, Liu L, Ju G, Zhang X, Xie L, Liu S, et al. An association study of the KPNB3 locus with schizophrenia in a Chinese population. Schizophr Res. 2005;76:363-5 pubmed
  76. Schaper S, Franke J, Meijsing S, Ehrenhofer Murray A. Nuclear import of the histone acetyltransferase complex SAS-I in Saccharomyces cerevisiae. J Cell Sci. 2005;118:1473-84 pubmed
    ..A database search based on the aligned consensus sequence revealed potential new import substrates of the Kap123p and Pse1p nuclear import pathways, which are connected to chromatin function. ..
  77. Hutchinson E, Orr O, Man Liu S, Engelhardt O, Fodor E. Characterization of the interaction between the influenza A virus polymerase subunit PB1 and the host nuclear import factor Ran-binding protein 5. J Gen Virol. 2011;92:1859-69 pubmed publisher
    ..Furthermore, mutations affecting RanBP5 binding are incompatible with or severely attenuate viral growth, providing further support for a key role for RanBP5 in the influenza A virus life cycle. ..
  78. De Iaco A, Luban J. Inhibition of HIV-1 infection by TNPO3 depletion is determined by capsid and detectable after viral cDNA enters the nucleus. Retrovirology. 2011;8:98 pubmed publisher
    ..TNPO3 promotes HIV-1 infectivity at a step in the virus life cycle that is detectable after the preintegration complex arrives in the nucleus and CA is the viral determinant for TNPO3 dependence. ..
  79. Freedman N, Yamamoto K. Importin 7 and importin alpha/importin beta are nuclear import receptors for the glucocorticoid receptor. Mol Biol Cell. 2004;15:2276-86 pubmed
    ..Interestingly, importin 7, importin 8, and importin alpha bound GR even in the absence of hormone; thus, hormonal control of localization is exerted at a step downstream of import receptor binding. ..
  80. Ciciarello M, Mangiacasale R, Thibier C, Guarguaglini G, Marchetti E, Di Fiore B, et al. Importin beta is transported to spindle poles during mitosis and regulates Ran-dependent spindle assembly factors in mammalian cells. J Cell Sci. 2004;117:6511-22 pubmed
    ..Targeting of TPX2/importin-beta complexes to poles is a key aspect in Ran-dependent control of the mitotic apparatus in mammalian cells. ..
  81. Kotera I, Sekimoto T, Miyamoto Y, Saiwaki T, Nagoshi E, Sakagami H, et al. Importin alpha transports CaMKIV to the nucleus without utilizing importin beta. EMBO J. 2005;24:942-51 pubmed
    ..This is the first report to demonstrate definitely that mammalian importin alpha solely carries a cargo protein into the nucleus without utilizing the classical importin beta-dependent transport system. ..
  82. Güttinger S, Mühlhäusser P, Koller Eichhorn R, Brennecke J, Kutay U. Transportin2 functions as importin and mediates nuclear import of HuR. Proc Natl Acad Sci U S A. 2004;101:2918-23 pubmed
    ..Finally, crosscompetition experiments demonstrated that HuR nucleocytoplasmic shuttling signal and M9 are imported along redundant pathways involving TRN1/2, substantiating the function of TRN2 in nuclear import. ..