PDBsum entry 1qgr

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protein Protein-protein interface(s) links
Transport receptor PDB id
Protein chains
872 a.a. *
24 a.a. *
Waters ×186
* Residue conservation analysis
PDB id:
Name: Transport receptor
Title: Structure of importin beta bound to the ibb domain of importin alpha (ii crystal form, grown at low ph)
Structure: Protein (importin beta subunit). Chain: a. Synonym: karyopherin beta-1, nuclear factor p97, importin b. Engineered: yes. Protein (importin alpha-2 subunit). Chain: b. Fragment: ibb domain. Synonym: importin alpha-2 subunit, karyopherin alpha-2
Source: Homo sapiens. Human. Organism_taxid: 9606. Cellular_location: cytoplasm. Expressed in: escherichia coli. Expression_system_taxid: 562. Synthetic: yes. Other_details: this protein is chemically synthesized. The sequence of this protein is naturally found in the
Biol. unit: Monomer (from PDB file)
2.30Å     R-factor:   0.235     R-free:   0.285
Authors: G.Cingolani,C.Petosa,K.Weis,C.W.Muller
Key ref:
G.Cingolani et al. (1999). Structure of importin-beta bound to the IBB domain of importin-alpha. Nature, 399, 221-229. PubMed id: 10353244 DOI: 10.1038/20367
04-May-99     Release date:   24-May-99    
Go to PROCHECK summary

Protein chain
Pfam   ArchSchema ?
Q14974  (IMB1_HUMAN) -  Importin subunit beta-1
876 a.a.
872 a.a.*
Protein chain
Pfam   ArchSchema ?
P52292  (IMA2_HUMAN) -  Importin subunit alpha-1
529 a.a.
24 a.a.
Key:    PfamA domain  Secondary structure  CATH domain
* PDB and UniProt seqs differ at 2 residue positions (black crosses)

 Gene Ontology (GO) functional annotation 
  GO annot!
  Cellular component     endoplasmic reticulum tubular network   11 terms 
  Biological process     viral reproduction   16 terms 
  Biochemical function     protein binding     8 terms  


DOI no: 10.1038/20367 Nature 399:221-229 (1999)
PubMed id: 10353244  
Structure of importin-beta bound to the IBB domain of importin-alpha.
G.Cingolani, C.Petosa, K.Weis, C.W.Müller.
Cytosolic proteins bearing a classical nuclear localization signal enter the nucleus bound to a heterodimer of importin-alpha and importin-beta (also called karyopherin-alpha and -beta). The formation of this heterodimer involves the importin-beta-binding (IBB) domain of importin-alpha, a highly basic amino-terminal region of roughly 40 amino-acid residues. Here we report the crystal structure of human importin-beta bound to the IBB domain of importin-alpha, determined at 2.5 A and 2.3 A resolution in two crystal forms. Importin-beta consists of 19 tandemly repeated HEAT motifs and wraps intimately around the IBB domain. The association involves two separate regions of importin-beta, recognizing structurally distinct parts of the IBB domain: an amino-terminal extended moiety and a carboxy-terminal helix. The structure indicates that significant conformational changes occur when importin-beta binds or releases the IBB domain domain and suggests how dissociation of the importin-alpha/beta heterodimer may be achieved upon nuclear entry.
  Selected figure(s)  
Figure 1.
Figure 1: Structure of importin- beta-bound to the IBB domain of importin- alpha-. a, View down the superhelical axis. A and B helices are shown in red and yellow, respectively, with connecting residues in grey. The acidic loop with the DDDDDW motif in HR8 is shown in blue. b, Side view. The ribbon representing importin- is coloured progressively from yellow to red as the chain proceeds from N to C terminus. The model has been rotated by 90° about a vertical axis with respect to a. c, Comparison of importin- and the protein phosphatase 2A PR65/A subunit. Proteins are represented as 'beads-on-a-string', with the centre of each HEAT repeat indicated by a sphere. d, Final 2Fo - Fc electron-density map (contoured at 1.0 ) of the N-terminal IBB moiety in crystal form I. Carbon atoms of the IBB domain and importin- are in green and yellow, respectively. Figs 1a–c, 4 and 5 were prepared with MOLSCRIPT^48.
Figure 2.
Figure 2: Primary and secondary structures of importin- beta-and the IBB domain. a, The IBB domain of human importin 2 aligned with eight homologues, including human snurportin1. Conserved residues are in red (acidic) or highlighted in blue (basic). Invariant residues are indicated by a black triangle, highly conserved hydrophobic residues by a colon, and Phe(Tyr)17 by a dot. b, Structural alignment of the 19 HEAT repeats. Conserved hydrophobic and polar residues are highlighted in green and red, respectively. The acidic loop is enclosed in a blue box. A line underneath residues indicates a helix, and a kink in the line marks a kink in the helix. c, A representative HEAT repeat. Consensus residue positions in HEAT-10 are shown. Leu(13) is actually Leu 413. Highly conserved residues Leu(13) and Ala(28) are boxed as in b.
  The above figures are reprinted by permission from Macmillan Publishers Ltd: Nature (1999, 399, 221-229) copyright 1999.  
  Figures were selected by the author.  

Literature references that cite this PDB file's key reference

  PubMed id Reference
21139563 M.Grünwald, and F.Bono (2011).
Structure of Importin13-Ubc9 complex: nuclear import and release of a key regulator of sumoylation.
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PDB code: 2xwu
21481773 T.L.Yeh, C.Y.Lee, L.M.Amzel, P.J.Espenshade, and M.A.Bianchet (2011).
The hypoxic regulator of sterol synthesis nro1 is a nuclear import adaptor.
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PDB code: 3msv
20023628 B.L.Sibanda, D.Y.Chirgadze, and T.L.Blundell (2010).
Crystal structure of DNA-PKcs reveals a large open-ring cradle comprised of HEAT repeats.
  Nature, 463, 118-121.
PDB code: 3kgv
20927102 E.H.Rubinson, A.S.Gowda, T.E.Spratt, B.Gold, and B.F.Eichman (2010).
An unprecedented nucleic acid capture mechanism for excision of DNA damage.
  Nature, 468, 406-411.
PDB codes: 3jx7 3jxy 3jxz 3jy1
19933700 I.S.Seong, J.M.Woda, J.J.Song, A.Lloret, P.D.Abeyrathne, C.J.Woo, G.Gregory, J.M.Lee, V.C.Wheeler, T.Walz, R.E.Kingston, J.F.Gusella, R.A.Conlon, and M.E.Macdonald (2010).
Huntingtin facilitates polycomb repressive complex 2.
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20826343 J.K.Forwood, A.Lange, U.Zachariae, M.Marfori, C.Preast, H.Grubmüller, M.Stewart, A.H.Corbett, and B.Kobe (2010).
Quantitative structural analysis of importin-β flexibility: paradigm for solenoid protein structures.
  Structure, 18, 1171-1183.
PDB code: 3nd2
21151903 T.J.Mosca, and T.L.Schwarz (2010).
Drosophila Importin-α2 is involved in synapse, axon and muscle development.
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19680239 A.G.Cook, N.Fukuhara, M.Jinek, and E.Conti (2009).
Structures of the tRNA export factor in the nuclear and cytosolic states.
  Nature, 461, 60-65.
PDB codes: 3ibv 3icq
19782947 B.D.Lazarus, D.C.Love, and J.A.Hanover (2009).
O-GlcNAc cycling: implications for neurodegenerative disorders.
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19965479 C.Okada, E.Yamashita, S.J.Lee, S.Shibata, J.Katahira, A.Nakagawa, Y.Yoneda, and T.Tsukihara (2009).
A High-Resolution Structure of the Pre-microRNA Nuclear Export Machinery.
  Science, 326, 1275-1279.
PDB code: 3a6p
19680228 D.Mohr, S.Frey, T.Fischer, T.Güttler, and D.Görlich (2009).
Characterisation of the passive permeability barrier of nuclear pore complexes.
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19421156 E.W.Debler, G.Blobel, and A.Hoelz (2009).
Nuclear transport comes full circle.
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19124778 J.C.Phillips (2009).
Scaling and self-organized criticality in proteins II.
  Proc Natl Acad Sci U S A, 106, 3113-3118.  
19204005 J.Kahle, E.Piaia, S.Neimanis, M.Meisterernst, and D.Doenecke (2009).
Regulation of nuclear import and export of negative cofactor 2.
  J Biol Chem, 284, 9382-9393.  
  20300488 K.Lui, and Y.Huang (2009).
RanGTPase: A Key Regulator of Nucleocytoplasmic Trafficking.
  Mol Cell Pharmacol, 1, 148-156.  
19801417 L.J.Terry, and S.R.Wente (2009).
Flexible gates: dynamic topologies and functions for FG nucleoporins in nucleocytoplasmic transport.
  Eukaryot Cell, 8, 1814-1827.  
19274657 R.Peters (2009).
Translocation through the nuclear pore: Kaps pave the way.
  Bioessays, 31, 466-477.  
19668212 S.M.Dias, K.F.Wilson, K.S.Rojas, A.L.Ambrosio, and R.A.Cerione (2009).
The molecular basis for the regulation of the cap-binding complex by the importins.
  Nat Struct Mol Biol, 16, 930-937.
PDB codes: 3fex 3fey
19389996 T.Monecke, T.Güttler, P.Neumann, A.Dickmanns, D.Görlich, and R.Ficner (2009).
Crystal structure of the nuclear export receptor CRM1 in complex with Snurportin1 and RanGTP.
  Science, 324, 1087-1091.
PDB code: 3gjx
19414022 Y.An, C.Y.Chen, B.Moyer, P.Rotkiewicz, M.A.Elsliger, A.Godzik, I.A.Wilson, and W.E.Balch (2009).
Structural and functional analysis of the globular head domain of p115 provides insight into membrane tethering.
  J Mol Biol, 391, 26-41.
PDB codes: 3gq2 3grl
18343812 A.Lange, R.E.Mills, S.E.Devine, and A.H.Corbett (2008).
A PY-NLS nuclear targeting signal is required for nuclear localization and function of the Saccharomyces cerevisiae mRNA-binding protein Hrp1.
  J Biol Chem, 283, 12926-12934.  
18187419 G.Mitrousis, A.S.Olia, N.Walker-Kopp, and G.Cingolani (2008).
Molecular basis for the recognition of snurportin 1 by importin beta.
  J Biol Chem, 283, 7877-7884.
PDB codes: 2p8q 2q5d
18606808 H.Sakiyama, R.M.Wynn, W.R.Lee, M.Fukasawa, H.Mizuguchi, K.H.Gardner, J.J.Repa, and K.Uyeda (2008).
Regulation of Nuclear Import/Export of Carbohydrate Response Element-binding Protein (ChREBP): INTERACTION OF AN {alpha}-HELIX OF ChREBP WITH THE 14-3-3 PROTEINS AND REGULATION BY PHOSPHORYLATION.
  J Biol Chem, 283, 24899-24908.  
18280242 H.Zheng, A.S.Olia, M.Gonen, S.Andrews, G.Cingolani, and T.Gonen (2008).
A conformational switch in bacteriophage p22 portal protein primes genome injection.
  Mol Cell, 29, 376-383.  
17682055 M.M.Pradeepa, S.Manjunatha, V.Sathish, S.Agrawal, and M.R.Rao (2008).
Involvement of importin-4 in the transport of transition protein 2 into the spermatid nucleus.
  Mol Cell Biol, 28, 4331-4341.  
19037259 N.Jin, C.Y.Chow, L.Liu, S.N.Zolov, R.Bronson, M.Davisson, J.L.Petersen, Y.Zhang, S.Park, J.E.Duex, D.Goldowitz, M.H.Meisler, and L.S.Weisman (2008).
VAC14 nucleates a protein complex essential for the acute interconversion of PI3P and PI(3,5)P(2) in yeast and mouse.
  EMBO J, 27, 3221-3234.  
18203617 S.Saint Fleur, and H.Fujii (2008).
Cytokine-induced nuclear translocation of signaling proteins and their analysis using the inducible translocation trap system.
  Cytokine, 41, 187-197.  
18547523 U.Zachariae, and H.Grubmüller (2008).
Importin-beta: structural and dynamic determinants of a molecular spring.
  Structure, 16, 906-915.  
18366719 V.A.Delmar, R.C.Chan, and D.J.Forbes (2008).
Xenopus importin beta validates human importin beta as a cell cycle negative regulator.
  BMC Cell Biol, 9, 14.  
17506639 A.Cook, F.Bono, M.Jinek, and E.Conti (2007).
Structural biology of nucleocytoplasmic transport.
  Annu Rev Biochem, 76, 647-671.  
17170104 A.Lange, R.E.Mills, C.J.Lange, M.Stewart, S.E.Devine, and A.H.Corbett (2007).
Classical nuclear localization signals: definition, function, and interaction with importin alpha.
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17056062 A.Paradise, M.K.Levin, G.Korza, and J.H.Carson (2007).
Significant proportions of nuclear transport proteins with reduced intracellular mobilities resolved by fluorescence correlation spectroscopy.
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18282135 A.V.Sorokin, E.R.Kim, and L.P.Ovchinnikov (2007).
Nucleocytoplasmic transport of proteins.
  Biochemistry (Mosc), 72, 1439-1457.  
17259172 D.Wohlwend, A.Strasser, A.Dickmanns, D.Doenecke, and R.Ficner (2007).
Thermodynamic analysis of H1 nuclear import: receptor tuning of importinbeta/importin7.
  J Biol Chem, 282, 10707-10719.  
17355870 J.Al-Bassam, N.A.Larsen, A.A.Hyman, and S.C.Harrison (2007).
Crystal structure of a TOG domain: conserved features of XMAP215/Dis1-family TOG domains and implications for tubulin binding.
  Structure, 15, 355-362.
PDB code: 2of3
19384428 J.E.Holt, J.D.Ly-Huynh, A.Efthymiadis, G.R.Hime, K.L.Loveland, and D.A.Jans (2007).
Regulation of Nuclear Import During Differentiation; The IMP alpha Gene Family and Spermatogenesis.
  Curr Genomics, 8, 323-334.  
17889670 K.C.Slep, and R.D.Vale (2007).
Structural basis of microtubule plus end tracking by XMAP215, CLIP-170, and EB1.
  Mol Cell, 27, 976-991.
PDB codes: 2qjx 2qjz 2qk0 2qk1 2qk2
17182855 K.J.Ryan, Y.Zhou, and S.R.Wente (2007).
The karyopherin Kap95 regulates nuclear pore complex assembly into intact nuclear envelopes in vivo.
  Mol Biol Cell, 18, 886-898.  
17596301 M.Frieman, B.Yount, M.Heise, S.A.Kopecky-Bromberg, P.Palese, and R.S.Baric (2007).
Severe acute respiratory syndrome coronavirus ORF6 antagonizes STAT1 function by sequestering nuclear import factors on the rough endoplasmic reticulum/Golgi membrane.
  J Virol, 81, 9812-9824.  
17537211 M.J.Pryor, S.M.Rawlinson, R.E.Butcher, C.L.Barton, T.A.Waterhouse, S.G.Vasudevan, P.G.Bardin, P.J.Wright, D.A.Jans, and A.D.Davidson (2007).
Nuclear localization of dengue virus nonstructural protein 5 through its importin alpha/beta-recognized nuclear localization sequences is integral to viral infection.
  Traffic, 8, 795-807.  
17287812 M.Stewart (2007).
Molecular mechanism of the nuclear protein import cycle.
  Nat Rev Mol Cell Biol, 8, 195-208.  
17308347 R.K.Nookala, S.Hussain, and L.Pellegrini (2007).
Insights into Fanconi Anaemia from the structure of human FANCE.
  Nucleic Acids Res, 35, 1638-1648.
PDB code: 2ilr
17938174 S.Neimanis, W.Albig, D.Doenecke, and J.Kahle (2007).
Sequence elements in both subunits of the DNA fragmentation factor are essential for its nuclear transport.
  J Biol Chem, 282, 35821-35830.  
17229728 T.Morita, A.Yamashita, I.Kashima, K.Ogata, S.Ishiura, and S.Ohno (2007).
Distant N- and C-terminal domains are required for intrinsic kinase activity of SMG-1, a critical component of nonsense-mediated mRNA decay.
  J Biol Chem, 282, 7799-7808.  
16522802 A.C.Hausrath, and A.Goriely (2006).
Repeat protein architectures predicted by a continuum representation of fold space.
  Protein Sci, 15, 753-760.  
16785238 A.E.Hodel, M.T.Harreman, K.F.Pulliam, M.E.Harben, J.S.Holmes, M.R.Hodel, K.M.Berland, and A.H.Corbett (2006).
Nuclear localization signal receptor affinity correlates with in vivo localization in Saccharomyces cerevisiae.
  J Biol Chem, 281, 23545-23556.  
16421734 A.S.Madrid, and K.Weis (2006).
Nuclear transport is becoming crystal clear.
  Chromosoma, 115, 98.  
16901787 B.J.Lee, A.E.Cansizoglu, K.E.Süel, T.H.Louis, Z.Zhang, and Y.M.Chook (2006).
Rules for nuclear localization sequence recognition by karyopherin beta 2.
  Cell, 126, 543-558.
PDB code: 2h4m
16902583 C.Rollenhagen, and N.Panté (2006).
Nuclear import of spliceosomal snRNPs.
  Can J Physiol Pharmacol, 84, 367-376.  
16567089 E.Conti, C.W.Müller, and M.Stewart (2006).
Karyopherin flexibility in nucleocytoplasmic transport.
  Curr Opin Struct Biol, 16, 237-244.  
16689644 H.Stark, and R.Lührmann (2006).
Cryo-electron microscopy of spliceosomal components.
  Annu Rev Biophys Biomol Struct, 35, 435-457.  
16240320 H.Yagisawa (2006).
Nucleocytoplasmic shuttling of phospholipase C-delta1: a link to Ca2+.
  J Cell Biochem, 97, 233-243.  
16698552 L.Bellsolell, P.F.Cho-Park, F.Poulin, N.Sonenberg, and S.K.Burley (2006).
Two structurally atypical HEAT domains in the C-terminal portion of human eIF4G support binding to eIF4A and Mnk1.
  Structure, 14, 913-923.  
16525863 M.A.van der Aa, E.Mastrobattista, R.S.Oosting, W.E.Hennink, G.A.Koning, and D.J.Crommelin (2006).
The nuclear pore complex: the gateway to successful nonviral gene delivery.
  Pharm Res, 23, 447-459.  
16407315 M.Arnold, A.Nath, D.Wohlwend, and R.H.Kehlenbach (2006).
Transportin is a major nuclear import receptor for c-Fos: a novel mode of cargo interaction.
  J Biol Chem, 281, 5492-5499.  
16704975 M.Arnold, A.Nath, J.Hauber, and R.H.Kehlenbach (2006).
Multiple importins function as nuclear transport receptors for the Rev protein of human immunodeficiency virus type 1.
  J Biol Chem, 281, 20883-20890.  
16868551 N.C.Reich, and L.Liu (2006).
Tracking STAT nuclear traffic.
  Nat Rev Immunol, 6, 602-612.  
17215878 R.Truant, R.Atwal, and A.Burtnik (2006).
Hypothesis: Huntingtin may function in membrane association and vesicular trafficking.
  Biochem Cell Biol, 84, 912-917.  
  16880557 T.Imasaki, T.Shimizu, H.Hashimoto, Y.Hidaka, M.Yamada, and M.Sato (2006).
Crystallization and preliminary X-ray crystallographic studies of transportin 1 in complex with nucleocytoplasmic shuttling and nuclear localization fragments.
  Acta Crystallogr Sect F Struct Biol Cryst Commun, 62, 785-787.  
16962977 U.Zachariae, and H.Grubmüller (2006).
A highly strained nuclear conformation of the exportin Cse1p revealed by molecular dynamics simulations.
  Structure, 14, 1469-1478.  
16595690 W.Li, L.C.Serpell, W.J.Carter, D.C.Rubinsztein, and J.A.Huntington (2006).
Expression and characterization of full-length human huntingtin, an elongated HEAT repeat protein.
  J Biol Chem, 281, 15916-15922.  
15907467 A.J.Stein, G.Fuchs, C.Fu, S.L.Wolin, and K.M.Reinisch (2005).
Structural insights into RNA quality control: the Ro autoantigen binds misfolded RNAs via its central cavity.
  Cell, 121, 529-539.
PDB codes: 1yvp 1yvr
15625192 C.F.Chuang, and C.I.Bargmann (2005).
A Toll-interleukin 1 repeat protein at the synapse specifies asymmetric odorant receptor expression via ASK1 MAPKKK signaling.
  Genes Dev, 19, 270-281.  
16030253 J.K.Ospina, G.B.Gonsalvez, J.Bednenko, E.Darzynkiewicz, L.Gerace, and A.G.Matera (2005).
Cross-talk between snurportin1 subdomains.
  Mol Biol Cell, 16, 4660-4671.  
16231302 J.Perry (2005).
Weighing in on a Timeless controversy.
  Proteins, 61, 699-703.  
15702987 L.F.Pemberton, and B.M.Paschal (2005).
Mechanisms of receptor-mediated nuclear import and nuclear export.
  Traffic, 6, 187-198.  
15919823 L.Liu, K.M.McBride, and N.C.Reich (2005).
STAT3 nuclear import is independent of tyrosine phosphorylation and mediated by importin-alpha3.
  Proc Natl Acad Sci U S A, 102, 8150-8155.  
15611084 M.H.Chen, I.Ben-Efraim, G.Mitrousis, N.Walker-Kopp, P.J.Sims, and G.Cingolani (2005).
Phospholipid scramblase 1 contains a nonclassical nuclear localization signal with unique binding site in importin alpha.
  J Biol Chem, 280, 10599-10606.
PDB code: 1y2a
15780942 M.M.Golas, B.Sander, C.L.Will, R.Lührmann, and H.Stark (2005).
Major conformational change in the complex SF3b upon integration into the spliceosomal U11/U12 di-snRNP as revealed by electron cryomicroscopy.
  Mol Cell, 17, 869-883.  
15864299 R.Nevo, V.Brumfeld, R.Kapon, P.Hinterdorfer, and Z.Reich (2005).
Direct measurement of protein energy landscape roughness.
  EMBO Rep, 6, 482-486.  
15864302 S.J.Lee, Y.Matsuura, S.M.Liu, and M.Stewart (2005).
Structural basis for nuclear import complex dissociation by RanGTP.
  Nature, 435, 693-696.
PDB code: 2bku
16338415 T.A.Isgro, and K.Schulten (2005).
Binding dynamics of isolated nucleoporin repeat regions to importin-beta.
  Structure, 13, 1869-1879.  
16222336 Y.Matsuura, and M.Stewart (2005).
Nup50/Npap60 function in nuclear protein import complex disassembly and importin recycling.
  EMBO J, 24, 3681-3689.
PDB codes: 2c1m 2c1t
15964794 Z.You, C.Chahwan, J.Bailis, T.Hunter, and P.Russell (2005).
ATM activation and its recruitment to damaged DNA require binding to the C terminus of Nbs1.
  Mol Cell Biol, 25, 5363-5379.  
15525506 A.Harel, and D.J.Forbes (2004).
Importin beta: conducting a much larger cellular symphony.
  Mol Cell, 16, 319-330.  
15574331 C.Petosa, G.Schoehn, P.Askjaer, U.Bauer, M.Moulin, U.Steuerwald, M.Soler-López, F.Baudin, I.W.Mattaj, and C.W.Müller (2004).
Architecture of CRM1/Exportin1 suggests how cooperativity is achieved during formation of a nuclear export complex.
  Mol Cell, 16, 761-775.
PDB code: 1w9c
15523559 D.Devos, S.Dokudovskaya, F.Alber, R.Williams, B.T.Chait, A.Sali, and M.P.Rout (2004).
Components of coated vesicles and nuclear pore complexes share a common molecular architecture.
  PLoS Biol, 2, e380.  
15350979 D.S.Goldfarb, A.H.Corbett, D.A.Mason, M.T.Harreman, and S.A.Adam (2004).
Importin alpha: a multipurpose nuclear-transport receptor.
  Trends Cell Biol, 14, 505-514.  
15545602 J.A.Roden, B.Belt, J.B.Ross, T.Tachibana, J.Vargas, and M.B.Mudgett (2004).
A genetic screen to isolate type III effectors translocated into pepper cells during Xanthomonas infection.
  Proc Natl Acad Sci U S A, 101, 16624-16629.  
14991001 L.Xu, and J.Massagué (2004).
Nucleocytoplasmic shuttling of signal transducers.
  Nat Rev Mol Cell Biol, 5, 209-219.  
14970380 M.Dlakić, and D.Tollervey (2004).
The Noc proteins involved in ribosome synthesis and export contain divergent HEAT repeats.
  RNA, 10, 351-354.  
15361863 M.Jínek, J.Rehwinkel, B.D.Lazarus, E.Izaurralde, J.A.Hanover, and E.Conti (2004).
The superhelical TPR-repeat domain of O-linked GlcNAc transferase exhibits structural similarities to importin alpha.
  Nat Struct Mol Biol, 11, 1001-1007.
PDB code: 1w3b
14729571 M.Oeffinger, M.Dlakic, and D.Tollervey (2004).
A pre-ribosome-associated HEAT-repeat protein is required for export of both ribosomal subunits.
  Genes Dev, 18, 196-209.  
14561738 N.Fukuhara, E.Fernandez, J.Ebert, E.Conti, and D.Svergun (2004).
Conformational variability of nucleo-cytoplasmic transport factors.
  J Biol Chem, 279, 2176-2181.  
15450977 N.Mosammaparast, and L.F.Pemberton (2004).
Karyopherins: from nuclear-transport mediators to nuclear-function regulators.
  Trends Cell Biol, 14, 547-556.  
15537541 S.J.Goldenberg, T.C.Cascio, S.D.Shumway, K.C.Garbutt, J.Liu, Y.Xiong, and N.Zheng (2004).
Structure of the Cand1-Cul1-Roc1 complex reveals regulatory mechanisms for the assembly of the multisubunit cullin-dependent ubiquitin ligases.
  Cell, 119, 517-528.
PDB code: 1u6g
15342649 W.Wang, X.Yang, T.Kawai, I.López de Silanes, K.Mazan-Mamczarz, P.Chen, Y.M.Chook, C.Quensel, M.Köhler, and M.Gorospe (2004).
AMP-activated protein kinase-regulated phosphorylation and acetylation of importin alpha1: involvement in the nuclear import of RNA-binding protein HuR.
  J Biol Chem, 279, 48376-48388.  
15602554 Y.Matsuura, and M.Stewart (2004).
Structural basis for the assembly of a nuclear export complex.
  Nature, 432, 872-877.
PDB code: 1wa5
12917401 B.Pyhtila, and M.Rexach (2003).
A gradient of affinity for the karyopherin Kap95p along the yeast nuclear pore complex.
  J Biol Chem, 278, 42699-42709.  
12594203 C.Koerner, T.Guan, L.Gerace, and G.Cingolani (2003).
Synergy of silent and hot spot mutations in importin beta reveals a dynamic mechanism for recognition of a nuclear localization signal.
  J Biol Chem, 278, 16216-16221.  
12802078 C.Rollenhagen, P.Mühlhäusser, U.Kutay, and N.Panté (2003).
Importin beta-depending nuclear import pathways: role of the adapter proteins in the docking and releasing steps.
  Mol Biol Cell, 14, 2104-2115.  
12604785 D.P.Denning, S.S.Patel, V.Uversky, A.L.Fink, and M.Rexach (2003).
Disorder in the nuclear pore complex: the FG repeat regions of nucleoporins are natively unfolded.
  Proc Natl Acad Sci U S A, 100, 2450-2455.  
12684370 I.Nikolaev, M.F.Cochet, and B.Felenbok (2003).
Nuclear import of zinc binuclear cluster proteins proceeds through multiple, overlapping transport pathways.
  Eukaryot Cell, 2, 209-221.  
12820967 J.B.Green, C.D.Gardner, R.P.Wharton, and A.K.Aggarwal (2003).
RNA recognition via the SAM domain of Smaug.
  Mol Cell, 11, 1537-1548.
PDB code: 1oxj
12885761 J.Bednenko, G.Cingolani, and L.Gerace (2003).
Importin beta contains a COOH-terminal nucleoporin binding region important for nuclear transport.
  J Cell Biol, 162, 391-401.  
12553904 J.Perry, and N.Kleckner (2003).
The ATRs, ATMs, and TORs are giant HEAT repeat proteins.
  Cell, 112, 151-155.  
12621039 K.Uchida, H.Miyauchi, T.Furuichi, T.Michikawa, and K.Mikoshiba (2003).
Critical regions for activation gating of the inositol 1,4,5-trisphosphate receptor.
  J Biol Chem, 278, 16551-16560.  
12600309 K.Weis (2003).
Regulating access to the genome: nucleocytoplasmic transport throughout the cell cycle.
  Cell, 112, 441-451.  
12917407 L.Xu, C.Alarcón, S.Cöl, and J.Massagué (2003).
Distinct domain utilization by Smad3 and Smad4 for nucleoporin interaction and nuclear import.
  J Biol Chem, 278, 42569-42577.  
12969505 M.Carrière, V.Escriou, A.Savarin, and D.Scherman (2003).
Coupling of importin beta binding peptide on plasmid DNA: transfection efficiency is increased by modification of lipoplex's physico-chemical properties.
  BMC Biotechnol, 3, 14.  
12738865 M.M.Golas, B.Sander, C.L.Will, R.Lührmann, and H.Stark (2003).
Molecular architecture of the multiprotein splicing factor SF3b.
  Science, 300, 980-984.  
14645832 M.Stewart (2003).
Structural biology. Nuclear trafficking.
  Science, 302, 1513-1514.  
12486120 M.T.Harreman, M.R.Hodel, P.Fanara, A.E.Hodel, and A.H.Corbett (2003).
The auto-inhibitory function of importin alpha is essential in vivo.
  J Biol Chem, 278, 5854-5863.  
12932731 P.Harjes, and E.E.Wanker (2003).
The hunt for huntingtin function: interaction partners tell many different stories.
  Trends Biochem Sci, 28, 425-433.  
12808444 R.Nevo, C.Stroh, F.Kienberger, D.Kaftan, V.Brumfeld, M.Elbaum, Z.Reich, and P.Hinterdorfer (2003).
A molecular switch between alternative conformational states in the complex of Ran and importin beta1.
  Nat Struct Biol, 10, 553-557.  
12571241 R.P.Darst, A.Dasgupta, C.Zhu, J.Y.Hsu, A.Vroom, T.Muldrow, and D.T.Auble (2003).
Mot1 regulates the DNA binding activity of free TATA-binding protein in an ATP-dependent manner.
  J Biol Chem, 278, 13216-13226.  
14501140 S.J.Lee, T.Sekimoto, E.Yamashita, E.Nagoshi, A.Nakagawa, H.Tanaka, Y.Yoneda, and T.Tsukihara (2003).
Crystallization and preliminary crystallographic analysis of the importin-beta-SREBP-2 complex.
  Acta Crystallogr D Biol Crystallogr, 59, 1866-1868.  
14645851 S.J.Lee, T.Sekimoto, E.Yamashita, E.Nagoshi, A.Nakagawa, N.Imamoto, M.Yoshimura, H.Sakai, K.T.Chong, T.Tsukihara, and Y.Yoneda (2003).
The structure of importin-beta bound to SREBP-2: nuclear import of a transcription factor.
  Science, 302, 1571-1575.
PDB code: 1ukl
12105224 A.J.Brooks, M.Johansson, A.V.John, Y.Xu, D.A.Jans, and S.G.Vasudevan (2002).
The interdomain region of dengue NS5 protein that binds to the viral helicase NS3 contains independently functional importin beta 1 and importin alpha/beta-recognized nuclear localization signals.
  J Biol Chem, 277, 36399-36407.  
12138188 A.R.Ball, J.A.Schmiesing, C.Zhou, H.C.Gregson, Y.Okada, T.Doi, and K.Yokomori (2002).
Identification of a chromosome-targeting domain in the human condensin subunit CNAP1/hCAP-D2/Eg7.
  Mol Cell Biol, 22, 5769-5781.  
12270919 A.V.Kajava (2002).
What curves alpha-solenoids? Evidence for an alpha-helical toroid structure of Rpn1 and Rpn2 proteins of the 26 S proteasome.
  J Biol Chem, 277, 49791-49798.  
12065587 D.P.Denning, V.Uversky, S.S.Patel, A.L.Fink, and M.Rexach (2002).
The Saccharomyces cerevisiae nucleoporin Nup2p is a natively unfolded protein.
  J Biol Chem, 277, 33447-33455.  
12504010 G.Cingolani, J.Bednenko, M.T.Gillespie, and L.Gerace (2002).
Molecular basis for the recognition of a nonclassical nuclear localization signal by importin beta.
  Mol Cell, 10, 1345-1353.
PDB code: 1m5n
12379151 H.Takano, and J.F.Gusella (2002).
The predominantly HEAT-like motif structure of huntingtin and its association and coincident nuclear entry with dorsal, an NF-kB/Rel/dorsal family transcription factor.
  BMC Neurosci, 3, 15.  
12442173 I.Bosanac, J.R.Alattia, T.K.Mal, J.Chan, S.Talarico, F.K.Tong, K.I.Tong, F.Yoshikawa, T.Furuichi, M.Iwai, T.Michikawa, K.Mikoshiba, and M.Ikura (2002).
Structure of the inositol 1,4,5-trisphosphate receptor binding core in complex with its ligand.
  Nature, 420, 696-700.
PDB code: 1n4k
11815630 J.Huber, A.Dickmanns, and R.Lührmann (2002).
The importin-beta binding domain of snurportin1 is responsible for the Ran- and energy-independent nuclear import of spliceosomal U snRNPs in vitro.
  J Cell Biol, 156, 467-479.  
12010460 J.T.Murray, C.Panaretou, H.Stenmark, M.Miaczynska, and J.M.Backer (2002).
Role of Rab5 in the recruitment of hVps34/p150 to the early endosome.
  Traffic, 3, 416-427.  
11875068 L.Shu, X.Zhang, and P.J.Houghton (2002).
Myogenic differentiation is dependent on both the kinase function and the N-terminal sequence of mammalian target of rapamycin.
  J Biol Chem, 277, 16726-16732.  
11854401 N.Panté, and M.Kann (2002).
Nuclear pore complex is able to transport macromolecules with diameters of about 39 nm.
  Mol Biol Cell, 13, 425-434.  
12372263 N.Vodovar, J.D.Clayton, R.Costa, M.Odell, and C.P.Kyriacou (2002).
The Drosophila clock protein Timeless is a member of the Arm/HEAT family.
  Curr Biol, 12, R610-R611.  
11923538 P.Kalab, K.Weis, and R.Heald (2002).
Visualization of a Ran-GTP gradient in interphase and mitotic Xenopus egg extracts.
  Science, 295, 2452-2456.  
12372823 R.Bayliss, T.Littlewood, L.A.Strawn, S.R.Wente, and M.Stewart (2002).
GLFG and FxFG nucleoporins bind to overlapping sites on importin-beta.
  J Biol Chem, 277, 50597-50606.
PDB codes: 1o6o 1o6p
12048190 R.Fagerlund, K.Mélen, L.Kinnunen, and I.Julkunen (2002).
Arginine/lysine-rich nuclear localization signals mediate interactions between dimeric STATs and importin alpha 5.
  J Biol Chem, 277, 30072-30078.  
12151228 S.Djordjevic, and P.C.Driscoll (2002).
Structural insight into substrate specificity and regulatory mechanisms of phosphoinositide 3-kinases.
  Trends Biochem Sci, 27, 426-432.  
12138183 S.Kuersten, G.J.Arts, T.C.Walther, L.Englmeier, and I.W.Mattaj (2002).
Steady-state nuclear localization of exportin-t involves RanGTP binding and two distinct nuclear pore complex interaction domains.
  Mol Cell Biol, 22, 5708-5720.  
  11423015 A.C.Ström, and K.Weis (2001).
Importin-beta-like nuclear transport receptors.
  Genome Biol, 2, REVIEWS3008.  
11709169 C.Steegborn, O.Danot, R.Huber, and T.Clausen (2001).
Crystal structure of transcription factor MalT domain III: a novel helix repeat fold implicated in regulated oligomerization.
  Structure, 9, 1051-1060.
PDB code: 1hz4
11229403 C.Wiese, A.Wilde, M.S.Moore, S.A.Adam, A.Merdes, and Y.Zheng (2001).
Role of importin-beta in coupling Ran to downstream targets in microtubule assembly.
  Science, 291, 653-656.  
11343901 E.Conti, and E.Izaurralde (2001).
Nucleocytoplasmic transport enters the atomic age.
  Curr Opin Cell Biol, 13, 310-319.  
11283257 E.Nagoshi, and Y.Yoneda (2001).
Dimerization of sterol regulatory element-binding protein 2 via the helix-loop-helix-leucine zipper domain is a prerequisite for its nuclear localization mediated by importin beta.
  Mol Cell Biol, 21, 2779-2789.  
11729264 I.G.Macara (2001).
Transport into and out of the nucleus.
  Microbiol Mol Biol Rev, 65, 570.  
11701921 I.R.Vetter, and A.Wittinghofer (2001).
The guanine nucleotide-binding switch in three dimensions.
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11172724 J.Marcotrigiano, I.B.Lomakin, N.Sonenberg, T.V.Pestova, C.U.Hellen, and S.K.Burley (2001).
A conserved HEAT domain within eIF4G directs assembly of the translation initiation machinery.
  Mol Cell, 7, 193-203.
PDB code: 1hu3
11571268 L.Englmeier, M.Fornerod, F.R.Bischoff, C.Petosa, I.W.Mattaj, and U.Kutay (2001).
RanBP3 influences interactions between CRM1 and its nuclear protein export substrates.
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11328871 L.Queimado, M.Rao, R.A.Schultz, E.V.Koonin, L.Aravind, T.Nardo, M.Stefanini, and E.C.Friedberg (2001).
Cloning the human and mouse MMS19 genes and functional complementation of a yeast mms19 deletion mutant.
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11416198 M.Sagermann, T.H.Stevens, and B.W.Matthews (2001).
Crystal structure of the regulatory subunit H of the V-type ATPase of Saccharomyces cerevisiae.
  Proc Natl Acad Sci U S A, 98, 7134-7139.
PDB code: 1ho8
  11251069 P.Maurer, M.Redd, J.Solsbacher, F.R.Bischoff, M.Greiner, A.V.Podtelejnikov, M.Mann, K.Stade, K.Weis, and G.Schlenstedt (2001).
The nuclear export receptor Xpo1p forms distinct complexes with NES transport substrates and the yeast Ran binding protein 1 (Yrb1p).
  Mol Biol Cell, 12, 539-549.  
11336677 T.A.Edwards, S.E.Pyle, R.P.Wharton, and A.K.Aggarwal (2001).
Structure of Pumilio reveals similarity between RNA and peptide binding motifs.
  Cell, 105, 281-289.
PDB code: 3h3d
11042144 A.F.Neuwald, and T.Hirano (2000).
HEAT repeats associated with condensins, cohesins, and other complexes involved in chromosome-related functions.
  Genome Res, 10, 1445-1452.  
10801418 A.J.Bannister, E.A.Miska, D.Görlich, and T.Kouzarides (2000).
Acetylation of importin-alpha nuclear import factors by CBP/p300.
  Curr Biol, 10, 467-470.  
10722750 C.Jabet, E.R.Sprague, A.P.VanDemark, and C.Wolberger (2000).
Characterization of the N-terminal domain of the yeast transcriptional repressor Tup1. Proposal for an association model of the repressor complex Tup1 x Ssn6.
  J Biol Chem, 275, 9011-9018.  
10842307 D.A.Jans, C.Y.Xiao, and M.H.Lam (2000).
Nuclear targeting signal recognition: a key control point in nuclear transport?
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10822175 D.Christophe, C.Christophe-Hobertus, and B.Pichon (2000).
Nuclear targeting of proteins: how many different signals?
  Cell Signal, 12, 337-341.  
10873829 D.J.Owen, and J.P.Luzio (2000).
Structural insights into clathrin-mediated endocytosis.
  Curr Opin Cell Biol, 12, 467-474.  
10745017 E.Conti, and J.Kuriyan (2000).
Crystallographic analysis of the specific yet versatile recognition of distinct nuclear localization signals by karyopherin alpha.
  Structure, 8, 329-338.
PDB codes: 1ee4 1ee5
10729155 E.Lombardo, J.C.Ramírez, M.Agbandje-McKenna, and J.M.Almendral (2000).
A beta-stranded motif drives capsid protein oligomers of the parvovirus minute virus of mice into the nucleus for viral assembly.
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11208133 J.A.Ybe, D.E.Wakeham, F.M.Brodsky, and P.K.Hwang (2000).
Molecular structures of proteins involved in vesicle fusion.
  Traffic, 1, 474-479.  
10791968 J.Hyman, H.Chen, P.P.Di Fiore, P.De Camilli, and A.T.Brunger (2000).
Epsin 1 undergoes nucleocytosolic shuttling and its eps15 interactor NH(2)-terminal homology (ENTH) domain, structurally similar to Armadillo and HEAT repeats, interacts with the transcription factor promyelocytic leukemia Zn(2)+ finger protein (PLZF).
  J Cell Biol, 149, 537-546.
PDB code: 1edu
10967423 J.K.Hood, and P.A.Silver (2000).
Diverse nuclear transport pathways regulate cell proliferation and oncogenesis.
  Biochim Biophys Acta, 1471, M31-M41.  
10713112 M.C.Lai, R.I.Lin, S.Y.Huang, C.W.Tsai, and W.Y.Tarn (2000).
A human importin-beta family protein, transportin-SR2, interacts with the phosphorylated RS domain of SR proteins.
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10851017 M.Sakaguchi, M.Miyazaki, Y.Inoue, T.Tsuji, H.Kouchi, T.Tanaka, H.Yamada, and M.Namba (2000).
Relationship between contact inhibition and intranuclear S100C of normal human fibroblasts.
  J Cell Biol, 149, 1193-1206.  
11129790 N.Imamoto (2000).
Diversity in nucleocytoplasmic transport pathways.
  Cell Struct Funct, 25, 207-216.  
11056478 P.S.Agutter, and D.N.Wheatley (2000).
Random walks and cell size.
  Bioessays, 22, 1018-1023.  
11208130 R.Bayliss, A.H.Corbett, and M.Stewart (2000).
The molecular mechanism of transport of macromolecules through nuclear pore complexes.
  Traffic, 1, 448-456.  
10929717 R.Bayliss, T.Littlewood, and M.Stewart (2000).
Structural basis for the interaction between FxFG nucleoporin repeats and importin-beta in nuclear trafficking.
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PDB code: 1f59
10666614 R.P.Baker, and M.Stewart (2000).
Crystallization and preliminary X-ray diffraction analysis of the Saccharomyces cerevisiae ran-binding protein Mog1p.
  Acta Crystallogr D Biol Crystallogr, 56, 229-231.  
11137006 S.Panizza, T.Tanaka, A.Hochwagen, F.Eisenhaber, and K.Nasmyth (2000).
Pds5 cooperates with cohesin in maintaining sister chromatid cohesion.
  Curr Biol, 10, 1557-1564.  
10827939 S.R.Wente (2000).
Gatekeepers of the nucleus.
  Science, 288, 1374-1377.  
10937870 T.Nishimoto (2000).
Upstream and downstream of ran GTPase.
  Biol Chem, 381, 397-405.  
  10888680 X.Pan, P.Roberts, Y.Chen, E.Kvam, N.Shulga, K.Huang, S.Lemmon, and D.S.Goldfarb (2000).
Nucleus-vacuole junctions in Saccharomyces cerevisiae are formed through the direct interaction of Vac8p with Nvj1p.
  Mol Biol Cell, 11, 2445-2457.  
10611974 D.Görlich, and U.Kutay (1999).
Transport between the cell nucleus and the cytoplasm.
  Annu Rev Cell Dev Biol, 15, 607-660.  
10375521 I.G.Macara (1999).
Nuclear transport: randy couples.
  Curr Biol, 9, R436-R439.  
10445030 L.M.Rice, and A.T.Brunger (1999).
Crystal structure of the vesicular transport protein Sec17: implications for SNAP function in SNARE complex disassembly.
  Mol Cell, 4, 85-95.
PDB code: 1qqe
10508790 S.J.Gamblin, and S.J.Smerdon (1999).
Nuclear transport: what a kary-on!
  Structure, 7, R199-R204.  
10490335 Y.M.Chook, G.Cingolani, E.Conti, M.Stewart, I.Vetter, and A.Wittinghofer (1999).
Pictures in cell biology. Structures of nuclear-transport components.
  Trends Cell Biol, 9, 310-311.  
The most recent references are shown first. Citation data come partly from CiteXplore and partly from an automated harvesting procedure. Note that this is likely to be only a partial list as not all journals are covered by either method. However, we are continually building up the citation data so more and more references will be included with time. Where a reference describes a PDB structure, the PDB code is shown on the right.