Nuclear envelope, nuclear pores, nucleocytoplasmic transport

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1 Nuclear envelope, nuclear pores, nucleocytoplasmic transport Know the organization of the nuclear envelope and associated proteins. Understand the organization of the nuclear pore complex. Understand the role of different types of nucleoporins. Be able to describe the nuclear import and nuclear export of proteins, and know the required components. Know different mechanisms by which nuclear import can be regulated. Know the basic principles of RNA nuclear export.

2 Nuclear envelope, nuclear pores, nucleocytoplasmic transport

3 Nuclear lamina (animal cell)

4 Foisner R J Cell Sci 2001;114:

5 SEVERAL MAJOR INTEGRAL MEMBRANE PROTEINS OF THE INNER NUCLEAR MEMBRANE INTERACT WITH BOTH THE NUCLEAR LAMINA AND CHROMATIN. The lamin B receptor (LBR), lamina-associated protein 2 (LAP2), Man-1, and emerin all bind lamin B. LBR associates with chromatin via HP1. The other three associate with chromatin via BAF. Emerin and LAP1 also bind to lamin A. Downloaded from: StudentConsult (on 16 December :26 AM) 2005 Elsevier

6 HUMAN DISEASES ASSOCIATED WITH NUCLEAR ENVELOPE ABNORMALITIES. Downloaded from: StudentConsult (on 16 December :26 AM) 2005 Elsevier

7 Page 704 Molecular Biology of the Cell ( Garland Science 2008)

8 Freeze fracture viewed with electron microscope Nuclear pores (NP) in nuclear envelope (NE)

9 Electron micrograph of a thin section showing face-on views of negatively stained nuclear pores. Figure 12-9(C) Figure 12-9c Molecular Biology of the Cell ( Garland Science 2008)

10 Nuclear pores as viewed by SEM, from the nuclear side of the nuclear envelope. This type of image clearly shows the nuclear basket. Xenopus oocytes nuclei have a very night density of nuclear pores, and are often used for these studies. Figure 12-9(B) Figure 12-9b Molecular Biology of the Cell ( Garland Science 2008)

11 Figure 12-9(A)

12 The molecular architecture of the nuclear pore complex. Alber F, Dokudovskaya S, Veenhoff LM, Zhang W, Kipper J, Devos D, Suprapto A, Karni-Schmidt O, Williams R, Chait BT, Sali A, Rout MP. Nature Nov 29;450(7170):

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16 The Outer Ring Complex Nups Form Solenoid- Beta Propeller Structures Devos et al. (2004) PLoS Biol 2(12): e380.

17 Outer Ring Nups are Similar to Vesicle Coat Proteins Devos et al. (2004) PLoS Biol 2(12): e380.

18 Model of nuclear pore evolucon Devos et al. (2004) PLoS Biol 2(12): e380.

19 GLE1 Nup42 Ndc1 Pom152 Pom34 Nuclear pores are similar across kingdoms Yeast Nup159 Nup82 Rae1 Nup166 Nup120 Nup133 Nup84 Seh1 Nup145C Sec13 Nup85 Nup192 Nup188 Nup157/170 Nic96 Nup53/59 Nup120 Nup133 Nup84 Seh1 Nup145C Sec13 Nup85 Rae1 Nup166 Nup2 Nsp1 Nup49 Nup57 Nup98 Rae1 Nup160 Nup43 Nup133 Nup107 Seh1 Nup96 Sec13 Nup75 Elys/HOS1 Nup62 Nup58 Nup54 Nup98 Rae1 Nup50 Nup205 Nup155 Nup93 Nup35 Nup160 Nup43 Nup133 Nup107 Seh1 Nup96 Sec13 Nup75 Elys/HOS1 Nup214 Nup88 CG1 gp210 NDC1 Nup136/ Nup1 Higher Plants Cytoplasm GLE1 Aladin RanGAP WIP GLE1 Aladin WIT CG1 Nup214 Nup88 gp210 NDC1 Rae1 Nup98 Nup160 Nup43 Nup133 Nup107 Seh1 Nup96 Sec13 Nup75 Elys/HOS1 Nup205 Nup155 Nup93 Nup35 Nup62 Nup58 Nup54 Nup160 Nup43 Nup133 Nup107 Seh1 Nup96 Sec13 Nup75 Elys/HOS1 Nup98 Rae1 Vertebrates RanGAP UBC9 Nup160 Nup43 Nup133 Nup37 Nup107 Seh1 Nup96 Sec13 Nup75 Elys Nup62 Nup58 Nup54 Nup45 Nup205 Nup188 Nup155 Nup93 Nup35 Nup358 Nup214 Nup88 Nup160 Nup43 Nup133 Nup37 Nup107 Seh1 Nup96 Sec13 Nup75 Elys CG1 gp210 Pom121 NDC1 Nup136/ Rae1 Nup98 Nup98 Rae1 Nup1 Nup153 Nup50 Nup50 GLE1 Aladin Nucleoplasm

20 Nuclear pores act as a diffusion barrier Molecules larger than ~50kDa cannot freely enter the nucleus Figure Figure Molecular Biology of the Cell ( Garland Science 2008)

21 A nuclear localization signal is necessary and sufficient for transport into the nucleus Figure 12-11

22 Nuclear Localization Signals Bind to an Import Receptor (or Importin ) Figure Figure Molecular Biology of the Cell ( Garland Science 2008)

23 Binding to Importins is Regulated by Ran Figure 12-16a Molecular Biology of the Cell ( Garland Science 2008)

24 Binding to Importins is Regulated by Ran Figure Figure 12-16b Molecular Biology of the Cell ( Garland Science 2008)

25 Figure 12-15

26 The Ran Cycle Impβ (or CAS) NTF2 Impβ (or CAS) NTF2 Figure Molecular Biology of the Cell ( Garland Science 2008)

27 It is, of course, more complicated Stewart (2007) Nature Reviews Molecular Cell Biology 8,

28 Nuclear Export Uses the Same Principles as Import Exportins are related to Importin β

29 Nuclear export of proteins

30 Regulation of nuclear transport

31 Regulated nuclear import is a step in signal transduction! Figure Molecular Biology of the Cell ( Garland Science 2008)

32 Figure Molecular Biology of the Cell ( Garland Science 2008)

33 Regulated nuclear import in plants Regulation of nucleocytoplasmic trafficking in plants.meier I, Somers DE.Curr Opin Plant Biol Oct;14(5):

34 Different RNAs exported from nucleus

35 Overview of the different RNA export pathways and the export factors

36 Transcription-coupled or splicing-coupled mrna export

37 Nuclear envelope, nuclear pores, nucleocytoplasmic transport Know the organization of the nuclear envelope and associated proteins. Understand the organization of the nuclear pore complex. Understand the role of different types of nucleoporins. Be able to describe the nuclear import and nuclear export of proteins, and know the required components. Know different mechanisms by which nuclear import can be regulated. Know the basic principles of RNA nuclear export.

Student Learning Outcomes: Nucleus distinguishes Eukaryotes from Prokaryotes

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