The dynamics of karyopherin-mediated nuclear transport

Biochemistry and Cell Biology

Volumn 79, Issue 5, 2001, Pages 603-612

  Marelli, Marcello ^a   Dilworth, David J ^a   Wozniak, Richard W ^b   Aitchison, John D ^b  

^a INSTITUTE FOR SYSTEMS BIOLOGY   (United States)

^b UNIVERSITY OF ALBERTA   (Canada)

Author keywords

Karyopherin; Nuclear pore complex; Nucleocytoplasmic transport; Nucleoporin; Nup2p

Indexed keywords

KARYOPHERIN BETA; NUCLEOPORIN; NUP2 PROTEIN, S CEREVISIAE; SACCHAROMYCES CEREVISIAE PROTEIN;

ACTIVE TRANSPORT; ANIMAL; BIOLOGICAL MODEL; CELL NUCLEUS MEMBRANE; CYTOPLASM; METABOLISM; PROTEIN TRANSPORT; REVIEW;

ACTIVE TRANSPORT, CELL NUCLEUS; ANIMALS; BETA KARYOPHERINS; CYTOPLASM; MODELS, BIOLOGICAL; NUCLEAR ENVELOPE; NUCLEAR PORE COMPLEX PROTEINS; PROTEIN TRANSPORT; SACCHAROMYCES CEREVISIAE PROTEINS;

EID: 85047683135     PISSN: 08298211     EISSN: None     Source Type: Journal    
DOI: 10.1139/o01-149     Document Type: Article

References (125)

1. Adam, E.J., and Adam, S.A. 1994. Identification of cytosolic factors required for nuclear location sequence-mediated binding to the nuclear envelope. J. Cell Biol. 125: 547-555.
2. Adam, S.A., and Gerace, L. 1991. Cytosolic proteins that specifically bind nuclear location signals are receptors for nuclear import. Cell, 66: 837-847.
3. Aitchison, J.D., Rout, M.P., Marelli, M., Blobel, G., and Wozniak, R.W. 1995. Two novel related yeast nucleoporins Nup170p and Nup157p: complementation with the vertebrate homologue Nup155p and functional interactions with the yeast nuclear pore-membrane protein Pom152p. J. Cell Biol. 131: 1133-1148.
4. Aitchison, J.D., Blobel, G., and Rout, M.P. 1996. Kap104p: a karyopherin involved in the nuclear transport of messenger RNA binding proteins. Science (Washington, D.C.), 274: 624-627.
5. Akey, C.W., and Goldfarb, D.S. 1989. Protein import through the nuclear pore complex is a multi-step process. J. Cell Biol. 109: 971-982.
6. Akey, C.W., and Radermacher, M. 1993. Architecture of the Xenopus nuclear pore complex revealed by three-dimensional cryo-electron microscopy. J. Cell Biol. 122: 1-19.
7. Arts, G.J., Kuersten, S., Romby, P., Ehresmann, B., and Mattaj, I.W. 1998. The role of exportin-t in selective nuclear export of mature tRNAs. EMBO J. 17: 7430-7441.
8. Bailer, S.M., Siniossoglou, S., Podtelejnikov, A., Hellwig, A., Mann, M., and Hurt, E. 1998. Nup116p and nup100p are interchangeable through a conserved motif which constitutes a docking site for the mRNA transport factor gle2p. EMBO J. 17: 1107-1119.
9. Bangs, P., Burke, B., Powers, C., Craig, R., Purohit, A., and Doxsey, S. 1998. Functional analysis of TPR: identification of nuclear pore complex association and nuclear localization domains and a role in mRNA export. J. Cell Biol. 143: 1801-1812.
10. Bastos, R., Lin, A., Enarson, M., and Burke, B. 1996. Targeting and function in mRNA export of nuclear pore complex protein Nup153. J. Cell Biol. 134: 1141-1156.
11. Bayliss, M.K., Little, D., Mallett, D.N., and Plumb, R.S. 2000. Crystallization and initial X-ray diffraction characterization of complexes of FxFG nucleoporin repeats with nuclear transport factors. J. Struct. Biol. 131: 240-247.
12. Ben-Efraim, I., and Gerace, L. 2001. Gradient of increasing affinity of importin beta for nucleoporins along the pathway of nuclear import. J. Cell Biol. 152: 411-418.
13. Bischoff, F.R., Krebber, H., Kempf, T., Hermes, I., and Ponstingl, H. 1995. Human RanGTPase-activating protein RanGAP1 is a homologue of yeast Rna1p involved in mRNA processing and transport. Proc. Natl. Acad. Sci. U.S.A. 92: 1749-1753.
14. Blobel, G. 1985. Gene gating: a hypothesis. Proc. Natl. Acad. Sci. U.S.A. 82: 8527-8529.
15. Booth, J.W., Belanger, K.D., Sannella, M.I., and Davis, L.I. 1999. The yeast nucleoporin Nup2p is involved in nuclear export of importin alpha/Srp1p. J. Biol. Chem. 274: 32 360 - 32 367.
16. Chi, N.C., Adam, E.J., and Adam, S.A. 1995. Sequence and characterization of cytoplasmic nuclear protein import factor p97. J. Cell Biol. 130: 265-274.
17. Cole, C.N., and Hammell, C.M. 1998. Nucleocytoplasmic transport: driving and directing transport. Curr. Biol. 8: R368-R372.
18. Cordes, V.C., Reidenbach, S., Rackwitz, H.R., and Franke, W.W. 1997. Identification of protein p270/Tpr as a constitutive component of the nuclear pore complex-attached intranuclear filaments. J. Cell Biol. 136: 531-544.
19. Dabauvalle, M.C., Benavente, R., and Chaly, N. 1988. Monoclonal antibodies to a Mr 68,000 pore complex glycoprotein interfere with nuclear protein uptake in Xenopus oocytes. Chromosoma, 97: 193-197.
20. Dilworth, D.J., Suprapto, A., Chait, B.T., Wozniak, R.W., Rout, M.P., and Aitchison, J.D. 2001. Nup2p dynamically associates with the distal regions of the yeaast nuclear pore complex. J. Cell Biol. 153: 1465-1478.
21. Dingwall, C., Kandels-Lewis, S., and Seraphin, B. 1995. A family of Ran binding proteins that includes nucleoporins. Proc. Natl. Acad. Sci. U.S.A. 92: 7525-7529.
22. Doye, V., and Hurt, E. 1997. From nucleoporins to nuclear pore complexes. Curr. Opin. Cell Biol. 9: 401-411.
23. Englmeier, L., Olivo, J.C., and Mattaj, I.W. 1999. Receptor-mediated substrate translocation through the nuclear pore complex without nucleotide triphosphate hydrolysis. Curr. Biol. 9: 30-41.
24. Fahrenkrog, B., Hurt, E.G., Aebi, U., and Pante, N. 1998. Molecular architecture of the yeast nuclear pore complex: localization of Nsp1p subcomplexes. J. Cell Biol. 143: 577-588.
25. Featherstone, C., Darby, M.K., and Gerace, L. 1988. A monoclonal antibody against the nuclear pore complex inhibits nucleocytoplasmic transport of protein and RNA in vivo. J. Cell Biol. 107: 1289-1297.
26. Feldherr, C.M., Kallenbach, E., and Schultz, N. 1984. Movement of a karyophilic protein through the nuclear pores of oocytes. Nippon Ganka Gakkai Zasshi, 88: 1548-1554.
27. Fontoura, B.M., Blobel, G., and Matunis, M.J. 1999. A conserved biogenesis pathway for nucleoporins: proteolytic processing of a 186-kilodalton precursor generates Nup98 and the novel nucleoporin, Nup96. J. Cell Biol. 144: 1097-1112.
28. Fontoura, B.M., Blobel, G., and Yaseen, N.R. 2000. The nucleoporin nup98 is a site for GDP/GTP exchange on ran and termination of karyopherin beta 2-mediated nuclear import. J. Biol. Chem. 275: 31 289-31 296.
29. Fornerod, M., Boer, J., van Baal, S., Jaegle, M., von Lindern, M., Murti, K.G., Davis, D., Bonten, J., Buijs, A., and Grosveld, G. 1995. Relocation of the carboxyl terminal part of CAN from the nuclear envelope to the nucleus as a result of leukemia-specific chromosome rearrangements. Oncogene, 10: 1739-1748.
30. Fornerod, M., Ohno, M., Yoshida, M., and Mattaj, I.W. 1997a. CRM1 is an export receptor for leucine-rich nuclear export signals. Cell, 90: 1051-1060.
31. Fornerod, M., van Deursen, J., van Baal, S., Reynolds, A., Davis, D., Murti, K.G., Fransen, J., and Grosveld, G. 1997b. The human homologue of yeast CRM1 is in a dynamic subcomplex with CAN/Nup214 and a novel nuclear pore component Nup88. EMBO J. 16: 807-816.
32. Fukuda, M., Asano, S., Nakamura, T., Adachi, M., Yoshida, M., Yanagida, M., and Nishida, E. 1997. CRM1 is responsible for intracellular transport mediated by the nuclear export signal. Nature (London), 390: 308-311.
33. Galy, V., Olivio-Marin, J.C., Schertan, H., Doye, V., Rascalou, N., and Nehrbass, U. 2000. Nuclear pore complexes in the organization of silent telomeric chromatin. Nature (London). 403: 108-112.
34. Görlich, D., and Kutay, U. 1999. Transport between the cell nucleus and the cytoplasm. Annu. Rev. Cell Dev. Biol. 15: 607-660.
35. Görlich, D., Prehn, S., Laskey, R.A., and Hartmann, E. 1994. Isolation of a protein that is essential for the first step of nuclear protein import. Cell, 79: 767-778.
36. Görlich, D., Kostka, S., Kraft, R., Dingwall, C., Laskey, R.A., Hartmann, E., and Prehn, S. 1995a. Two different subunits of importin cooperate to recognize nuclear localization signals and bind them to the nuclear envelope. Curr. Biol. 5: 383-392.
37. Görlich, D., Vogel, F., Mills, A.D., Hartmann, E., and Laskey, R.A. 1995b. Distinct functions for the two importin subunits in nuclear protein import. Nature (London), 377: 246-248.
38. Görlich, D., Pante, N., Kutay, U., Aebi, U., and Bischoff, F.R. 1996a. Identification of different roles for RanGDP and RanGTP in nuclear protein import. EMBO J. 15: 5584-5594.
39. Görlich, D., Kraft, R., Kostka, S., Vogel, F., Hartmann, E., Laskey, R.A., Mattaj, I.W., and Izaurraide, E. 1996b. Importin provides a link between nuclear protein import and U snRNA export. Cell, 87: 21-32.
40. Ho, A.K., Raczniak, G.A., Ives, E.B., and Wente, S.R. 2000. The integral membrane protein snl1p is genetically linked to yeast nuclear pore complex function Mol. Biol. Cell, 9: 355-373.
41. Hood, J.K., and Silver, P.A. 1999. In or out? regulating nuclear transport. Curr. Opin. Cell Biol. 11: 241-247.
42. Imamoto, N., Shimamoto, T., Kose, S., Takao, T., Tachibana, T., Matsubae, M., Sekimoto, T., Shimonishi, Y., and Yoneda, Y. 1995a. The nuclear pore-targeting complex binds to nuclear pores after association with a karyophile. FEBS Lett. 368: 415-419.
43. Imamoto, N., Tachibana, T., Matsubae, M., and Yoneda, Y. 1995b. A karyophilic protein forms a stable complex with cytoplasmic components prior to nuclear pore binding. J. Biol. Chem. 270: 8559-8565.
44. Iovine, M.K., and Wente, S.R. 1997. A nuclear export signal in Kap95p is required for both recycling the import factor and interaction with the nucleoporin GLFG repeat regions of Nup116p and Nup100p. J. Cell Biol. 137: 797-811.
45. Izaurralde, E., Lewis, J., Gamberi, C., Jarmolowski, A., McGuigan, C., and Mattaj, I.W. 1995. A cap-binding protein complex mediating U snRNA export. Nature (London), 376: 709-712.
46. Jarmolowski, A., Boelens, W.C., Izaurralde, E., and Mattaj, I.W. 1994. Nuclear export of different classes of RNA is mediated by specific factors. J. Cell Biol. 124: 627-635.
47. Kaffman, A., Rank, N.M., O'Shea, E.K. 1998a. Phosphorylation regulates association of the transcription factor Pho4 with its import receptor Pse1/Kap121. Genes Dev. 12: 2673-2683.
48. Kaffman, A., Rank, N.M., O'Neill, E.M., Huang, L.S., and O'Shea, E.K. 19986. The receptor Msn5 exports the phosphorylated transcription factor Pho4 out of the nucleus. Nature (London), 396: 482-486.
49. Kehlenbach, R.H., Dickmanns, A., Kehlenbach, A., Guan, T., and Gerace, L. 1999. A role for RanBP1 in the release of CRM1 from the nuclear pore complex in a terminal step of nuclear export. J. Cell Biol. 145: 645-657.
50. Kosova, B., Pante, N., Rollenhagen, C., Podtelejnikov, A., Mann, M., Aebi, U., and Hurt, E. 2000. Mlp2p, a component of nuclear pore attached intranuclear filaments, associates with nic96p. J. Biol. Chem. 275: 343-350.
51. Kraemer, D., and Blobel, G. 1997. mRNA binding protein mrnp 41 localizes to both nucleus and cytoplasm. Proc. Natl. Acad. Sci. U.S.A. 94: 9119-9124.
52. Kraemer, D., Wozniak, R.W., Blobel, G., and Radu, A. 1994. The human CAN protein, a putative oncogene product associated with myeloid leukemogenesis, is a nuclear pore complex protein that faces the cytoplasm. Proc. Natl. Acad. Sci. U.S.A. 91: 1519-1523.
53. Kraemer, D.M., Strambio-de-Castillia, C., Blobel, G., and Rout, M.P. 1995. The essential yeast nucleoporin NUP159 is located on the cytoplasmic side of the nuclear pore complex and serves in karyopherin-mediated binding of transport substrate. J. Biol. Chem. 270: 19017 - 19021.
54. Kutay, U., Bischoff, F.R., Kostka, S., Kraft, R., and Görlich, D. 1997. Export of importin alpha from the nucleus is mediated by a specific nuclear transport factor. Cell, 90: 1061-1071.
55. Kutay, U., Lipowsky, G., Izaurralde, E., Bischoff, F.R., Schwarzmaier, P., Hartmann, E., and Görlich, D. 1998. Identification of a tRNA-specific nuclear export receptor. Mol. Cell, 1: 359-369.
56. Lee, D.C., and Aitchison, J.D. 1999. Kap104p-mediated nuclear import. Nuclear localization signals in mRNA-binding proteins and the role of Ran and Rna. J. Biol. Chem. 274: 29031-29037.
57. Loeb, J.D., Davis, L.I., and Fink, G.R. 1993. NUP2, a novel yeast nucleoporin, has functional overlap with other proteins of the nuclear pore complex. Mol. Biol. Cell, 4: 209-222.
58. Mahajan, R., Gerace, L., and Melchior, F. 1998. Molecular characterization of the SUMO-1 modification of RanGAP1 and its role in nuclear envelope association. J. Cell Biol. 140: 259-270.
59. Marelli, M., Aitchison, J.D., and Wozniak, R.W. 1998. Specific binding of the karyopherin kap121p to a subunit of the nuclear pore complex containing nup53p, nup59p, and nup170p. J. Cell Biol. 143: 1813-1830.
60. Mattaj, I.W., and Englmeier, L. 1998. Nucleocytoplasmic transport: the soluble phase. J. Cell Biol. 143: 23-34.
61. Matunis, M.J., Wu, J., and Blobel, G. 1998. SUMO-1 modification and its role in targeting the ran GTPase-activating protein, RanGAP1, to the nuclear pore complex. J. Cell Biol. 140: 499-509.
62. Maurer, P., Redd, M., Solsbacher, J., Bischoff, F.R., Greiner, M., Podtelejnikov, A.V., Mann, M., Stade, K., Weis, K., and Schlenstedt, G. 2001. The nuclear export receptor xpo1p forms distinct complexes with nes transport substrates and the yeast ran binding protein 1 (yrb1). Mol. Biol. Cell, 12: 539-549.
63. Melchior, F., and Gerace, L. 1998. Two-way trafficking with Ran. Trends Cell Biol. 8: 175-179.
64. Melchior, F., Paschal, B., Evans, J., and Gerace, L. 1993. Inhibition of nuclear protein import by nonhydrolyzable analogues of GTP and identification of the small GTPase Ran/TC4 as an essential transport factor. J. Cell Biol. 123: 1649-1659.
65. Moore, M.S., and Blobel, G. 1993. The GTP-binding protein Ran/TC4 is required for protein import into the nucleus. Nature (London), 365: 661-663.
66. Moore, M.S., and Blobel, G. 1994. Purification of a Raninteracting protein that is required for protein import into the nucleus. Proc. Natl. Acad. Sci. U.S.A. 91: 10 212 - 10 216.
67. Moroianu, J., Hijikata, M., Blobel, G., and Radu, A. 1995. Mammalian karyopherin alpha 1 beta and alpha 2 beta heterodimers: alpha 1 or alpha 2 subunit binds nuclear localization signal and beta subunit interacts with peptide repeat-containing nucleoporins. Proc. Natl. Acad. Sci. U.S.A. 92: 6532-6536.
68. Moroianu, J., Blobel, G., and Radu, A. 1997. RanGTP-mediated nuclear export of karyopherin alpha involves its interaction with the nucleoporin Nup153. Proc. Natl. Acad. Sci. U.S.A. 94: 9699-9704.
69. Mueller, L., Cordes, V.C., Bischoff, F.R., and Ponstingl, H. 1998. Human RanBP3, a group of nuclear RanGTP binding proteins. FEBS Lett. 427: 330-336.
70. Nakielny, S., and Dreyfuss, G. 1999. Transport of proteins and RNAs in and out of the nucleus. Cell, 99: 677-690.
71. Nakielny, S., Shaikh, S., Burke, B., and Dreyfuss, G. 1999. Nup153 is an M9-containing mobile nucleoporin with a novel Ran-binding domain. EMBO J. 18: 1982-1995.
72. Nehrbass, U., and Blobel, G. 1996. Role of the nuclear transport factor p10 in nuclear import. Science (Washington, D.C.), 272: 120-122.
73. Nehrbass, U., Rout, M.P., Maguire, S., Blobel, G., and Wozniak, R.W. 1996. The yeast nucleoporin Nup188p interacts genetically and physically with the core structures of the nuclear pore complex. J. Cell Biol. 133: 1153-1162.
74. Neville, M., Stutz, F., Lee, L., Davis, L.I., and Rosbash, M. 1997. The importin-beta family member Crm1p bridges the interaction between Rev and the nuclear pore complex during nuclear export. Curr. Biol. 7: 767-775.
75. Ossareh-Nazari, B., Bachelerie, F., and Dargemont, C. 1997. Evidence for a role of CRM1 in signal-mediated nuclear protein export. Science (Washington, D.C.), 278: 141-144.
76. Paddy, M.R. 1998. The Tpr protein: linking structure and function in the nuclear interior? Am. J. Hum. Genet. 63: 305-310.
77. Pante, N., and Aebi, U. 1996. Sequential binding of import ligands to distinct nucleopore regions during their nuclear import. Science (Washington, D.C.), 273: 1729-1732.
78. Paschal, B.M., and Gerace, L. 1995. Identification of NTF2, a cytosolic factor for nuclear import that interacts with nuclear pore complex protein p62. J. Cell Biol. 129: 925-937.
79. Pemberton, L.F., Blobel, G., and Rosenblum, J.S. 1998. Transport routes through the nuclear pore complex. Curr. Opin. Cell Biol. 10: 392-399.
80. Percipalle, P., Clarkson, W.D., Kent, H.M., Rhodes, D., and Stewart, M. 1997. Molecular interactions between the importin alpha/beta heterodimer and proteins involved in vertebrate nuclear protein import. J. Mol. Biol. 266: 722-732.
81. Powers, M.A., Macaulay, C., Masiarz, F.R., and Forbes, D.J. 1995. Reconstituted nuclei depleted of a vertebrate GLFG nuclear pore protein, p97, import but are defective in nuclear growth and replication. J. Cell Biol. 128: 721-736.
82. Powers, M.A., Forbes, D.J., Dahlberg, J.E., and Lund, E. 1997. The vertebrate GLFG nucleoporin, Nup98, is an essential component of multiple RNA export pathways. J. Cell Biol. 136: 241-250.
83. Radu, A., Blobel, G., and Moore, M.S. 1995a. Identification of a protein complex that is required for nuclear protein import and mediates docking of import substrate to distinct nucleoporins. Proc. Natl. Acad. Sci. U.S.A. 92: 1769-1773.
84. Radu, A., Moore, M.S., and Blobel, G. 1995b. The peptide repeat domain of nucleoporin Nup98 functions as a docking site in transport across the nuclear pore complex. Cell, 81: 215-222.
85. Reichelt, R., Holzenburg, A., Buhle, E.L., Jr., Jarnik, M., Engel, A., Aebi, U. 1990. Correlation between structure and mass distribution of the nuclear pore complex and of distinct pore complex components. J. Cell Biol. 110: 883-894.
86. Rexach, M., and Blobel, G. 1995. Protein import into nuclei: association and dissociation reactions involving transport substrate, transport factors, and nucleoporins. Cell, 83: 683-692.
87. Ribbeck, K., and Görlich, D. 2001. Kinetic analysis of translocation through nuclear pore complexes. EMBO J. 20: 1320-1330.
88. Ribbeck, K., Lipowsky, G., Kent, H.M., Stewart, M., and Görlich, D. 1998. NTF2 mediates nuclear import of Ran. EMBO J. 17: 6587-6598.
89. Ribbeck, K., Kutay, U., Paraskeva, E., and Gorlich, D. 1999. The translocation of transportin-cargo complexes through nuclear pores is independent of both Ran and energy. Curr. Biol. 9: 47-50.
90. Richardson, W.D., Mills, A.D., Dilworth, S.M., Laskey, R.A., and Dingwall, C. 1988. Nuclear protein migration involves two steps: rapid binding at the nuclear envelope followed by slower translocation through nuclear pores. Cell, 52: 655-664.
91. Rout, M.P., and Aitchison, J.D. 2001. The nuclear pore complex as a transport machine. J. Biol. Chem. 276: 16 593 - 16 596.
92. Rout, M.P., and Blobel, G. 1993. Isolation of the yeast nuclear pore complex. J. Cell Biol. 123: 771-783.
93. Rout, M.P., and Wente, S.R. 1994. Pores for thought: nuclear pore proteins. Trends Cell Biol. 4: 357-365.
94. Rout, M.P., Blobel, G., and Aitchison, J.D. 1997. A distinct nuclear import pathway used by ribosomal proteins. Cell, 89: 715-725.
95. Rout, M.P., Aitchison, J.D., Suprapto, A., Hjertaas, K., Zhao, Y., and Chait, B.T. 2000. The yeast nuclear pore complex: composition, architecture, and transport mechanism. J. Cell Biol. 148: 635-651.
96. Sarkar, S., and Hopper, A.K. 1998. tRNA nuclear export in Saccharomyces cerevisiae: In situ hybridization analysis. Mol. Biol. Cell, 9: 3041-3055.
97. Schlenstedt, G., Wong, D.H., Koepp, D.M., and Silver, P.A. 1995. Mutants in a yeast ran binding protein are defective in nuclear transport. EMBO J. 14: 5367-5378.
98. Schlenstedt, G., Smirnova, E., Deane, R., Solsbacher, J., Kutay, U., Görlich, D., Ponstingl, H., and Bischoff, F.R. 1997. Yrb4p, a yeast ran-GTP-binding protein involved in import of ribosomal protein L25 into the nucleus. EMBO J. 16: 6237-6249.
99. Schwoebel, E.D., Talcott, B., Cushman, I., and Moore, M.S. 1998. Ran-dependent signal-mediated nuclear import does not require GTP hydrolysis by Ran. J. Biol. Chem. 273: 35 170 - 35 175.
100. Seedorf, M., and Silver, P.A. 1997. Importin/karyopherin protein family members required for mRNA export from the nucleus. Proc. Natl. Acad. Sci. U.S.A. 94: 8590-8595.
101. Seedorf, M., Damelin, M., Kahana, J., Taura, T., and Silver, P.A. 1999. Interactions between a nuclear transporter and a subset of nuclear pore complex proteins depend on Ran GTPase. Mol. Cell. Biol. 19: 1547-1557.
102. Shah, S., and Forbes, D.J. 1998. Separate nuclear import pathways converge on the nucleoporin Nup153 and can be dissected with dominant-negative inhibitors. Curr. Biol. 318: 1376-1386.
103. Shah, S., Tugendreich, S., and Forbes, D. 1998. Major binding sites for the nuclear import receptor are the internal nucleoporin Nup153 and the adjacent nuclear filament protein Tpr. J. Cell Biol. 141: 31-49.
104. Smith, A., Brownawell, A., and Macara, I.G. 1998. Nuclear import of Ran is mediated by the transport factor NTF2. Curr. Biol. 318: 1403-1406.
105. Solsbacher, J., Maurer, P., Bischoff, F.R., and Schlenstedt, G. 1998. Cse1p is involved in export of yeast importin alpha from the nucleus. Mol. Cell. Biol. 18: 6805-6815.
106. Solsbacher, J., Maurer, P., Vogel, F., and Schlenstedt, G. 2000. Nup2p, a yeast nucleoporin, functions in bidirectional transport of importin alpha. Mol. Cell. Biol. 20: 8468-8479.
107. Stade, K., Ford, C.S., Guthrie, C., and Weis, K. 1997. Exportin 1 (Crm1p) is an essential nuclear export factor. Cell, 90: 1041-1050.
108. Stoffler, D., Fahrenkrog, B., and Aebi, U. 1999. The nuclear pore complex: from molecular architecture to functional dynamics. Curr. Opin. Cell Biol. 11: 391-401.
109. Strambio-de-Castillia, C., Blobel, G., and Rout, M.P. 1999. Proteins connecting the nuclear pore complex with the nuclear interior. J. Cell Biol. 144: 839-855.
110. Sukegawa, J., and Blobel, G. 1993. A nuclear pore complex protein that contains zinc finger motifs, binds DNA, and faces the nucleoplasm. Cell, 72: 29-38.
111. Taura, T., Krebber, H., and Silver, P.A. 1998. A member of the Ran-binding protein family, Yrb2p, is involved in nuclear protein export. Proc. Natl. Acad. Sci. U.S.A. 95: 7427-7432.
112. Traglia, H.M., O'Connor, J.P., Tung, K.S., Dallabrida, S., Shen, W.C., and Hopper, A.K. 1996. Nucleus-associated pools of Rna1, the Saccharomyces cerevisiae Ran/TC4 GTPase activating protein involved in nucleus/cytosol transit. Proc. Natl. Acad. Sci. U.S.A. 93: 7667-7672.
113. Uetz, P., Giot, L., Cagney, G., Mansfield, T.A., Judson, R.S., Knight, J.R., Lockshon, D., Narayan, V., Srinivasan, M., Pochart, P., Qureshi-Emili, A., Li, Y., Godwin, B., Conover, D., Kalbfleisch, T., Vijayadamodar, G., Yang, M., Johnston, M., Fields, S., and Rothberg, J.M. 2000. A comprehensive analysis of protein-protein interactions in Saccharomyces cerevisiae. Nature (London), 403: 623-627.
114. van Deursen, J., Boer, J., Kasper, L., and Grosveld, G. 1996. G2 arrest and impaired nucleocytoplasmic transport in mouse embryos lacking the proto-oncogene CAN/Nup214. EMBO J. 15: 5574-5583.
115. Weis, K., Mattaj, I.W., and Lamond, A.I. 1995. Identification of hSRP1 alpha as a functional receptor for nuclear localization sequences. Science (Washington, D.C.), 268: 1049-1053.
116. Wente, S.R. 2000. Gatekeepers of the nucleus. Science (Washington, D.C.), 288: 1374-1377.
117. Wozniak, R.W., Rout, M.P., and Aitchison, J.D. 1998. Karyopherins and kissing cousins. Trends Cell Biol. 8: 184-188.
118. Wu, J., Matunis, M.J., Kraemer, D., Blobel, G., and Coutavas, E. 1995. Nup358, a cytoplasmically exposed nucleoporin with peptide repeats, Ran-GTP binding sites, zinc fingers, a cyclophilin A homologous domain, and a leucine-rich region. J. Biol. Chem. 270: 14209 - 14213.
119. Yang, Q., Rout, M.P., and Akey, C.W. 1998. Three-dimensional architecture of the isolated yeast nuclear pore complex: functional and evolutionary implications. Mol. Cell, 1: 519-529.
120. Yaseen, N.R., and Blobel, G. 1997. Cloning and characterization of human karyopherin beta3. Proc. Natl. Acad. Sci. U.S.A. 94: 4451-4456.
121. Yaseen, N.R., and Blobel, G. 1999. Two distinct classes of Ranbinding sites on the nucleoporin Nup-358. Proc. Natl. Acad. Sci. U.S.A. 96: 5516-5521.
122. Yokoyama, N., Hayashi, N., Seki, T., Pante, N., Ohba, T., Nishii, K., Kuma, K., Hayashida, T., Miyata, T., and Aebi, U., 1995. A giant nucleopore protein that binds Ran/TC4. Nature (London), 376: 184-188.
123. Yoshida, K., and Blobel, G. 2001. The karyopherin Kap142/Msn5p mediates nuclear import and nuclear export of different cargo proteins. J. Cell Biol. 152: 729-740.
124. Zabel, U., Doye, V., Tekotte, H., Wepf, R., Grandi, P., and Hurt, E.C. 1996. Nic96p is required for nuclear pore formation and functionally interacts with a novel nucleoporin, Nup188p. J. Cell Biol. 133: 1141-1152.
125. Zimowska, G., Ans, J.P., and Paddy, M.R. 1997. A Drosophila Tpr protein homolog is localized both in the extrachromosomal channel network and to nuclear pore complexes. Kaibogaku Zasshi, 72: 123-133.