The Nuclear Pore Complex as a Transport Machine

nuclear envelope nuclear pore complex nucleoporin ribonucleoprotein nuclear localization signal Nucleocytoplasmic Transport and the Nuclear Pore ComplexUnlike their prokaryotic counterparts, eukaryotic cells separate the nuclear synthesis of DNA and RNA from cytoplasmic protein synthesis with a barrier termed the nuclear envelope (NE).1 The NE is perforated by large proteinaceous assemblies, called nuclear pore complexes (NPCs), which act as the sole gatekeepers controlling the exchange of material between the two locales (reviewed in Ref. 1Wente S.R. Science. 2000; 288: 1374-1377Crossref PubMed Scopus (215) Google Scholar). NPCs are freely permeable to small molecules (such as water and ions), but they restrict the movement of larger molecules (such as proteins and RNAs) across the NE. To overcome this barrier, macromolecules carry specific signals that allow them to access the nucleocytoplasmic transport machinery of the cell. In this way the cell ensures that only selected macromolecules can travel between the nucleus and cytoplasm (reviewed in Ref. 2Mattaj I.W. Englmeier L. Annu. Rev. Biochem. 1998; 67: 265-306Crossref PubMed Scopus (1002) Google Scholar).Operationally, NPCs are composed of proteins called nucleoporins (or Nups) forming the stationary phase for nucleocytoplasmic exchange, whereas the mobile phase consists of soluble transport factors and their cargoes. As nucleocytoplasmic transport is driven by a series of specific interactions between components of both phases, it is frequently difficult to determine which proteins are permanent constituents of the NPC. Nevertheless, to understand how transport occurs, we must characterize the players in both phases and understand how their interplay leads to the coordinated vectorial exchange of macromolecules across the NE. In this review we focus on recent results that shed light on how some of these proteins interact to contribute to the elaborate NPC architecture and its function as a transport machine.Architecture of the NPCNPCs from different organisms share a common fundamental architecture (3Yang Q. Rout M.P. Akey C.W. Mol. Cell. 1998; 1: 223-234Abstract Full Text Full Text PDF PubMed Scopus (281) Google Scholar). These similarities likely provide clues as to the key features common to a functioning transport machine. The NPC is a large octagonally symmetric cylindrical structure. In yeast it estimated to be ∼50 MDa, whereas in metazoans it is over twice this mass (3Yang Q. Rout M.P. Akey C.W. Mol. Cell. 1998; 1: 223-234Abstract Full Text Full Text PDF PubMed Scopus (281) Google Scholar, 4Reichelt R. Holzenburg A. Buhle Jr., E.L. Jarnik M. Engel A. Aebi U. J. PubMed Scopus Google Scholar). that a it be that the NPC of different it that the yeast NPC only different proteins and the NPC a M.P. A. J. 2000; PubMed Scopus Google Scholar). the of how a large complex can be from The to in the of the The NPC is of a cylindrical from which the nucleus and cytoplasm (reviewed in Ref. J. 2000; PubMed Google The of the NPC is in the only is it composed of but is both in a to the NE and in a the cylindrical As from this nucleoporins are in and and are to both the nuclear and cytoplasmic of the NE M.P. A. J. 2000; PubMed Scopus Google Scholar, J. 2000; PubMed Google Scholar, Aebi U. PubMed Scopus Google this with the large of nucleoporins between and it how the NPC can be from a small of the large of nucleoporins them to between of the NPC. for we by NPC in and how they to the of NPC of a NPC from the cytoplasm the cytoplasmic NPC from the the nuclear The of the are by protein by and of both and yeast The of the and are on both the and of the NPC. The of the cytoplasmic nuclear and are on a the of the the of the NPC. by and and Akey of the NPC to the of the nucleoporins the whereas mass is of the to the interactions between nucleoporins M.P. A. J. 2000; PubMed Scopus Google Scholar, J. M. 2000; PubMed Scopus Google Pore and of the Nuclear nuclear envelope is composed of The NE is with the and the the Nuclear are by a of these two the the pore The the with the and proteins to this are termed is the function of they likely a in NPC by the of the pore and as a for the NPC. between from different but as the of pore is it likely that and to be J. PubMed Scopus Google Scholar, M. Jr., Mol. Cell. PubMed Scopus Google the of the pore must be and with the of the that the the this a pore but and are by pore proteins as which is in the 1998; PubMed Scopus Google Scholar, R. M. J. PubMed Google Scholar). a pore protein to be a of the is in NPC R. M. J. PubMed Google Scholar). is the of and this be to the of the NPC and nuclear envelope PubMed Scopus Google Scholar). is a protein in the of in yeast a for in NPC as is NE yeast The and of the NPC is to be the that and the of the NPC. As be the yeast nucleoporins that to this are to both of the NPC M.P. A. J. 2000; PubMed Scopus Google Scholar, Aebi U. PubMed Scopus Google Scholar). only of the nucleoporins is in is likely a of the and of the that proteins this with and contribute to that is to the of is by and of the for this is the M. Aebi U. J. 2000; PubMed Scopus Google Scholar). proteins in this complex interactions with of their a of protein interactions the J. M. 2000; PubMed Scopus Google Scholar). only two of the complex are in yeast this that the complex can of components by the complex a and a mass of the are NPC and in estimated M.P. A. J. 2000; PubMed Scopus Google Scholar, M. Aebi U. J. 2000; PubMed Scopus Google Scholar). this for of the yeast this complex to the of the NPC it that the of the to of the of the NPC M. Aebi U. J. 2000; PubMed Scopus Google of the in and yeast to the of in a and elaborate nuclear and cytoplasmic (3Yang Q. Rout M.P. Akey C.W. Mol. Cell. 1998; 1: 223-234Abstract Full Text Full Text PDF PubMed Scopus (281) Google Scholar, Aebi U. PubMed Scopus Google Scholar, C.W. M. J. PubMed Scopus Google Scholar). the features of the and complex are between the as they are in the for is can its the yeast protein Rout M.P. M. J. PubMed Scopus Google Scholar). The yeast that of this and a complex can be some of the yeast complex J. PubMed Scopus Google the contribute to NPC As transport across the NE the the of the the is to the must the of the NPC as a barrier to to in the nuclear the NPC and large the complex and the in different by C.W. M. J. PubMed Scopus Google Scholar, C.W. J. Full Text PDF PubMed Scopus Google Scholar). These different a of the from a permeable to only molecules to the of a by the of the transport cargoes. the and nuclear the transport of large as ribonucleoprotein J. Mol. PubMed Scopus Google Scholar, M. Akey J. 1998; PubMed Google in of the components and the of the NPC R. J. 2000; PubMed Scopus Google Scholar). These results are the to nucleoporins in controlling the NPC and that these proteins are of the proteins that Transport and the of the the These nucleoporins are to the mobile phase of transport and a in with and their cargoes. for the nucleus carry a nuclear localization signal whereas to be from the nucleus nuclear (reviewed in Ref. 2Mattaj I.W. Englmeier L. Annu. Rev. Biochem. 1998; 67: 265-306Crossref PubMed Scopus (1002) Google Scholar). The signals in by a of soluble transport proteins termed as and (reviewed in 2Mattaj I.W. Englmeier L. Annu. Rev. Biochem. 1998; 67: 265-306Crossref PubMed Scopus (1002) Google and Rout M.P. 1998; Full Text Full Text PDF PubMed Scopus Google Scholar). Transport as nuclear and small nuclear nuclear that are by their transport is by the small and 1998; Full Text Full Text PDF PubMed Google and J. 1998; Full Text Full Text PDF PubMed Scopus Google Scholar). that the complex the cytoplasmic and the NPC C.W. J. PubMed Scopus Google Scholar, Cell. Full Text PDF PubMed Scopus Google Scholar). it is that nuclear is by a series of and as the complex nucleoporins from the cytoplasmic of the NPC the to the the complex is to the nuclear PubMed Scopus Google of Transport the to understand how the NPC contribute to it is to characterize its The NPC is with nucleoporins by the of the These are in the nucleoporins and the of by of These nucleoporins to be the and are the from the of the cytoplasmic the to the of the nuclear M.P. A. J. 2000; PubMed Scopus Google Scholar, Aebi U. PubMed Scopus Google Scholar). As the nucleoporins are to be to the mobile phase and interact with of the as transport S.R. 2000; PubMed Scopus Google they are in movement across the of the of to a that likely interact with the of the in the the with structure. nucleoporins to M. J. PubMed Google and with the of the NPC (reviewed in J. 2000; PubMed Google and Aebi U. PubMed Scopus Google Scholar). is with these proteins forming the of the that from the and the and be As be from their from the of the in nucleoporins are to the by the of their M. R. 1998; PubMed Scopus Google Scholar, A. L. J. PubMed Google Scholar, J. PubMed Scopus Google Scholar, M. A. Aebi U. Mol. Cell. PubMed Scopus Google Transport transport can nucleoporins that to of transport factors (reviewed in Ref. S.R. 2000; PubMed Scopus Google Scholar). and the that of some to the NPC can be to that the NPC in the of the NPC and the of nucleoporins be transport NPC. this a of for transport for a of nucleoporins (reviewed in Ref. S.R. 2000; PubMed Scopus Google Scholar). allow different to different a the between different and the transport in both it that a NPC is of both and different transport J. PubMed Scopus Google Scholar). The NPC as a way to by a nucleoporin to a nuclear transport of the of this is the between and in in that with different nucleoporins the both that is a specific for M. J. 1998; PubMed Scopus Google Scholar, M. Mol. Cell. 2000; Full Text Full Text PDF PubMed Scopus Google Scholar). it that over the is and is a in the of the to the NPC it to be this results in a specific cell in nuclear M. J. 1998; PubMed Scopus Google of the and the of the nucleoporins in it is that nucleoporins are the whereas nucleoporins to the cytoplasmic are from the that the of transport factors the NPC is by the nucleoporins the of the NPC. likely that are of the of the NPC in which and transport factors as they the the complexes by the nucleoporins are both the yeast nucleoporin and the complex are on both of the NPC the M.P. A. J. 2000; PubMed Scopus Google Scholar, M. Aebi U. L. Mol. Cell. PubMed Scopus Google Scholar, M. U. R. R. J. PubMed Google Scholar, Aebi U. J. 1998; PubMed Scopus Google Scholar). with the that is only in the between from different Annu. Rev. PubMed Scopus Google that the in the of the and in their the NPCs of different of to nucleocytoplasmic exchange The only of this is the small (reviewed in 1998; Full Text Full Text PDF PubMed Google and J. 1998; Full Text Full Text PDF PubMed Scopus Google Scholar). as the is to it is likely that the from a across the NPC by the of of in the nucleus and in the cytoplasm transport by the and of transport complexes in the their they interact with in the whereas to their cargoes. the on is is the in the can their but between its and on its The different of to transport by and their cargoes. of for is soluble in both the and it is to the NPC two different protein The is to the cytoplasmic protein both and and in the cytoplasmic nucleoporin J. J. Full Text Full Text PDF PubMed Scopus Google Scholar). The of by a and is on both U. A. PubMed Scopus Google and the nucleoporin J. PubMed Scopus Google Scholar). to these nuclear and cytoplasmic components of the NPC a of in the of the nuclear the of the transport the exchange of between transport factors and the across the NPC. In the of to the NPC R. L. Cell. Full Text Full Text PDF PubMed Scopus (1002) Google Scholar, J. J. 1998; PubMed Scopus Google Scholar). and to the nucleoporin provide a of and of transport factors and their be in the large cells soluble factors the to from the In this it is that in the yeast the only protein to with the NPC U. A. PubMed Scopus Google is that the of the NPC be for of ribonucleoprotein complexes to the NPC with their nucleoporin the RNA is with both yeast and its the protein is to the NPC (reviewed in Ref. S.R. 2000; PubMed Scopus Google Scholar). These proteins in the in the to it the in the of the provide for the of the across the of must be to a for transport the NPC. the NPC a barrier to the of macromolecules across the but it must be to the of macromolecules the As to be to the of the the and of the be by NPC The is that the of the and the of the nucleoporins likely across the NPC transport across the NPC is by a of transport factors that interact with the large of that the of the NPC can be in the of transport by the with their to the nucleoporins As of these nucleoporins are on both of the transport factors exchange between nucleoporins on both of the NE. The NPC is a as proteins that can the NPC the barrier of the freely that is a composed of the M.P. A. J. 2000; PubMed Scopus Google Scholar). The is that the nuclear and cytoplasmic of the NPC likely contribute with the to the of transport M.P. A. J. 2000; PubMed Scopus Google Scholar). of that a nucleoporin to the transport of complexes U. Aebi U. J. PubMed Scopus Google Scholar, M. J. Full Text Full Text PDF PubMed Scopus Google Scholar, A. A. L. J. PubMed Scopus Google and only a and two it to determine the of the interactions and of the mobile and stationary phase components and to to understand in how these two phases to transport across the nuclear we must and to nucleocytoplasmic transport of to the of cells the of eukaryotic Nucleocytoplasmic Transport and the Nuclear Pore ComplexUnlike their prokaryotic counterparts, eukaryotic cells separate the nuclear synthesis of DNA and RNA from cytoplasmic protein synthesis with a barrier termed the nuclear envelope (NE).1 The NE is perforated by large proteinaceous assemblies, called nuclear pore complexes (NPCs), which act as the sole gatekeepers controlling the exchange of material between the two locales (reviewed in Ref. 1Wente S.R. Science. 2000; 288: 1374-1377Crossref PubMed Scopus (215) Google Scholar). NPCs are freely permeable to small molecules (such as water and ions), but they restrict the movement of larger molecules (such as proteins and RNAs) across the NE. To overcome this barrier, macromolecules carry specific signals that allow them to access the nucleocytoplasmic transport machinery of the cell. In this way the cell ensures that only selected macromolecules can travel between the nucleus and cytoplasm (reviewed in Ref. 2Mattaj I.W. Englmeier L. Annu. Rev. Biochem. 1998; 67: 265-306Crossref PubMed Scopus (1002) Google Scholar).Operationally, NPCs are composed of proteins called nucleoporins (or Nups) forming the stationary phase for nucleocytoplasmic exchange, whereas the mobile phase consists of soluble transport factors and their cargoes. As nucleocytoplasmic transport is driven by a series of specific interactions between components of both phases, it is frequently difficult to determine which proteins are permanent constituents of the NPC. Nevertheless, to understand how transport occurs, we must characterize the players in both phases and understand how their interplay leads to the coordinated vectorial exchange of macromolecules across the NE. In this review we focus on recent results that shed light on how some of these proteins interact to contribute to the elaborate NPC architecture and its function as a transport machine. their prokaryotic counterparts, eukaryotic cells separate the nuclear synthesis of DNA and RNA from cytoplasmic protein synthesis with a barrier termed the nuclear envelope (NE).1 The NE is perforated by large proteinaceous assemblies, called nuclear pore complexes (NPCs), which act as the sole gatekeepers controlling the exchange of material between the two locales (reviewed in Ref. 1Wente S.R. Science. 2000; 288: 1374-1377Crossref PubMed Scopus (215) Google Scholar). NPCs are freely permeable to small molecules (such as water and ions), but they restrict the movement of larger molecules (such as proteins and RNAs) across the NE. To overcome this barrier, macromolecules carry specific signals that allow them to access the nucleocytoplasmic transport machinery of the cell. In this way the cell ensures that only selected macromolecules can travel between the nucleus and cytoplasm (reviewed in Ref. 2Mattaj I.W. Englmeier L. Annu. Rev. Biochem. 1998; 67: 265-306Crossref PubMed Scopus (1002) Google Scholar). NPCs are composed of proteins called nucleoporins (or Nups) forming the stationary phase for nucleocytoplasmic exchange, whereas the mobile phase consists of soluble transport factors and their cargoes. As nucleocytoplasmic transport is driven by a series of specific interactions between components of both phases, it is frequently difficult to determine which proteins are permanent constituents of the NPC. Nevertheless, to understand how transport occurs, we must characterize the players in both phases and understand how their interplay leads to the coordinated vectorial exchange of macromolecules across the NE. In this review we focus on recent results that shed light on how some of these proteins interact to contribute to the elaborate NPC architecture and its function as a transport machine. of the NPCNPCs from different organisms share a common fundamental architecture (3Yang Q. Rout M.P. Akey C.W. Mol. Cell. 1998; 1: 223-234Abstract Full Text Full Text PDF PubMed Scopus (281) Google Scholar). These similarities likely provide clues as to the key features common to a functioning transport machine. The NPC is a large octagonally symmetric cylindrical structure. In yeast it estimated to be ∼50 MDa, whereas in metazoans it is over twice this mass (3Yang Q. Rout M.P. Akey C.W. Mol. Cell. 1998; 1: 223-234Abstract Full Text Full Text PDF PubMed Scopus (281) Google Scholar, 4Reichelt R. Holzenburg A. Buhle Jr., E.L. Jarnik M. Engel A. Aebi U. J. PubMed Scopus Google Scholar). that a it be that the NPC of different it that the yeast NPC only different proteins and the NPC a M.P. A. J. 2000; PubMed Scopus Google Scholar). the of how a large complex can be from The to in the of the The NPC is of a cylindrical from which the nucleus and cytoplasm (reviewed in Ref. J. 2000; PubMed Google The of the NPC is in the only is it composed of but is both in a to the NE and in a the cylindrical As from this nucleoporins are in and and are to both the nuclear and cytoplasmic of the NE M.P. A. J. 2000; PubMed Scopus Google Scholar, J. 2000; PubMed Google Scholar, Aebi U. PubMed Scopus Google this with the large of nucleoporins between and it how the NPC can be from a small of the large of nucleoporins them to between of the NPC. for we by NPC in and how they to the of the NPC to the of the nucleoporins the whereas mass is of the to the interactions between nucleoporins M.P. A. J. 2000; PubMed Scopus Google Scholar, J. M. 2000; PubMed Scopus Google NPCs from different organisms share a common fundamental architecture (3Yang Q. Rout M.P. Akey C.W. Mol. Cell. 1998; 1: 223-234Abstract Full Text Full Text PDF PubMed Scopus (281) Google Scholar). These similarities likely provide clues as to the key features common to a functioning transport machine. The NPC is a large octagonally symmetric cylindrical structure. In yeast it estimated to be ∼50 MDa, whereas in metazoans it is over twice this mass (3Yang Q. Rout M.P. Akey C.W. Mol. Cell. 1998; 1: 223-234Abstract Full Text Full Text PDF PubMed Scopus (281) Google Scholar, 4Reichelt R. Holzenburg A. Buhle Jr., E.L. Jarnik M. Engel A. Aebi U. J. PubMed Scopus Google Scholar). that a it be that the NPC of different it that the yeast NPC only different proteins and the NPC a M.P. A. J. 2000; PubMed Scopus Google Scholar). the of how a large complex can be from The to in the of the The NPC is of a cylindrical from which the nucleus and cytoplasm (reviewed in Ref. J. 2000; PubMed Google The of the NPC is in the only is it composed of but is both in a to the NE and in a the cylindrical As from this nucleoporins are in and and are to both the nuclear and cytoplasmic of the NE M.P. A. J. 2000; PubMed Scopus Google Scholar, J. 2000; PubMed Google Scholar, Aebi U. PubMed Scopus Google this with the large of nucleoporins between and it how the NPC can be from a small of the large of nucleoporins them to between of the NPC. for we by NPC in and how they to the machine. The Pore and of the Nuclear nuclear envelope is composed of The NE is with the and the the Nuclear are by a of these two the the pore The the with the and proteins to this are termed is the function of they likely a in NPC by the of the pore and as a for the NPC. between from different but as the of pore is it likely that and to be J. PubMed Scopus Google Scholar, M. Jr., Mol. Cell. PubMed Scopus Google the of the pore must be and with the of the that the the this a pore but and are by pore proteins as which is in the 1998; PubMed Scopus Google Scholar, R. M. J. PubMed Google Scholar). a pore protein to be a of the is in NPC R. M. J. PubMed Google Scholar). is the of and this be to the of the NPC and nuclear envelope PubMed Scopus Google Scholar). is a protein in the of in yeast a for in NPC as is NE yeast The nuclear envelope is composed of The NE is with the and the the Nuclear are by a of these two the the pore The the with the and proteins to this are termed is the function of they likely a in NPC by the of the pore and as a for the NPC. between from different but as the of pore is it likely that and to be NPCs J. PubMed Scopus Google Scholar, M. Jr., Mol. Cell. PubMed Scopus Google the of the pore must be and with the of the that the the this a pore but and are by pore proteins as which is in the 1998; PubMed Scopus Google Scholar, R. M. J. PubMed Google Scholar). a pore protein to be a of the is in NPC R. M. J. PubMed Google Scholar). is the of and this be to the of the NPC and nuclear envelope PubMed Scopus Google Scholar). is a protein in the of in yeast a for in NPC as is NE yeast The The and of the NPC is to be the that and the of the NPC. As be the yeast nucleoporins that to this are to both of the NPC M.P. A. J. 2000; PubMed Scopus Google Scholar, Aebi U. PubMed Scopus Google Scholar). only of the nucleoporins is in is likely a of the and of the that proteins this with and contribute to that is to the of is by and of the for this is the M. Aebi U. J. 2000; PubMed Scopus Google Scholar). proteins in this complex interactions with of their a of protein interactions the J. M. 2000; PubMed Scopus Google Scholar). only two of the complex are in yeast this that the complex can of components by the complex a and a mass of the are NPC and in estimated M.P. A. J. 2000; PubMed Scopus Google Scholar, M. Aebi U. J. 2000; PubMed Scopus Google Scholar). this for of the yeast this complex to the of the NPC it that the of the to of the of the NPC M. Aebi U. J. 2000; PubMed Scopus Google of the in and yeast to the of in a and elaborate nuclear and cytoplasmic (3Yang Q. Rout M.P. Akey C.W. Mol. Cell. 1998; 1: 223-234Abstract Full Text Full Text PDF PubMed Scopus (281) Google Scholar, Aebi U. PubMed Scopus Google Scholar, C.W. M. J. PubMed Scopus Google Scholar). the features of the and complex are between the as they are in the for is can its the yeast protein Rout M.P. M. J. PubMed Scopus Google Scholar). The yeast that of this and a complex can be some of the yeast complex J. PubMed Scopus Google the contribute to NPC As transport across the NE the the of the the is to the must the of the NPC as a barrier to to in the nuclear the NPC and large the complex and the in different by C.W. M. J. PubMed Scopus Google Scholar, C.W. J. Full Text PDF PubMed Scopus Google Scholar). These different a of the from a permeable to only molecules to the of a by the of the transport cargoes. the and nuclear the transport of large as ribonucleoprotein J. Mol. PubMed Scopus Google Scholar, M. Akey J. 1998; PubMed Google in of the components and the of the NPC R. J. 2000; PubMed Scopus Google Scholar). These results are the to nucleoporins in controlling the NPC and that these proteins are of the proteins that The of the NPC is to be the that and the of the NPC. As be the yeast nucleoporins that to this are to both of the NPC M.P. A. J. 2000; PubMed Scopus Google Scholar, Aebi U. PubMed Scopus Google Scholar). only of the nucleoporins is in is likely a of the and of the that proteins this with and contribute to that is to the of is by and of the for this is the M. Aebi U. J. 2000; PubMed Scopus Google Scholar). proteins in this complex interactions with of their a of protein interactions the J. M. 2000; PubMed Scopus Google Scholar). only two of the complex are in yeast this that the complex can of components by the complex a and a mass of the are NPC and in estimated M.P. A. J. 2000; PubMed Scopus Google Scholar, M. Aebi U. J. 2000; PubMed Scopus Google Scholar). this for of the yeast this complex to the of the NPC it that the of the to of the of the NPC M. Aebi U. J. 2000; PubMed Scopus Google Scholar). of the in and yeast to the of in a and elaborate nuclear and cytoplasmic (3Yang Q. Rout M.P. Akey C.W. Mol. Cell. 1998; 1: 223-234Abstract Full Text Full Text PDF PubMed Scopus (281) Google Scholar, Aebi U. PubMed Scopus Google Scholar, C.W. M. J. PubMed Scopus Google Scholar). the features of the and complex are between the as they are in the for is can its the yeast protein Rout M.P. M. J. PubMed Scopus Google Scholar). The yeast that of this and a complex can be some of the yeast complex J. PubMed Scopus Google Scholar). the contribute to NPC As transport across the NE the the of the the is to the must the of the NPC as a barrier to to in the nuclear the NPC and large the complex and the in different by C.W. M. J. PubMed Scopus Google Scholar, C.W. J. Full Text PDF PubMed Scopus Google Scholar). These different a of the from a permeable to only molecules to the of a by the of the transport cargoes. the and nuclear the transport of large as ribonucleoprotein J. Mol. PubMed Scopus Google Scholar, M. Akey J. 1998; PubMed Google Scholar). in of the components and the of the NPC R. J. 2000; PubMed Scopus Google Scholar). These results are the to nucleoporins in controlling the NPC and that these proteins are of the proteins that Nucleocytoplasmic Transport and the of the the These nucleoporins are to the mobile phase of transport and a in with and their cargoes. for the nucleus carry a nuclear localization signal whereas to be from the nucleus nuclear (reviewed in Ref. 2Mattaj I.W. Englmeier L. Annu. Rev. Biochem. 1998; 67: 265-306Crossref PubMed Scopus (1002) Google Scholar). The signals in by a of soluble transport proteins termed as and (reviewed in 2Mattaj I.W. Englmeier L. Annu. Rev. Biochem. 1998; 67: 265-306Crossref PubMed Scopus (1002) Google and Rout M.P. 1998; Full Text Full Text PDF PubMed Scopus Google Scholar). Transport as nuclear and small nuclear nuclear that are by their transport is by the small and 1998; Full Text Full Text PDF PubMed Google and J. 1998; Full Text Full Text PDF PubMed Scopus Google Scholar). that the complex the cytoplasmic and the NPC C.W. J. PubMed Scopus Google Scholar, Cell. Full Text PDF PubMed Scopus Google Scholar). it is that nuclear is by a series of and as the complex nucleoporins from the cytoplasmic of the NPC the to the the complex is to the nuclear PubMed Scopus

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