Function. 8 The Nucleus 9 Protein Sorting and Transport 10 Bioenergetics and Metabolism 11 The Cytoskeleton and Cell Movement 12 The Cell Surface
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1 PART III Cell Structure and Function 8 The Nucleus 9 Protein Sorting and Transport 10 Bioenergetics and Metabolism 11 The Cytoskeleton and Cell Movement 12 The Cell Surface 8 The Nucleus The Nuclear Envelope and Traffic Between the Nucleus and Cytoplasm Internal Organization of the Nucleus The Nucleolus The Nucleus During Mitosis 1
2 The Nuclear Envelope and Traffic Between the Nucleus and Cytoplasm Structure of the Nuclear Envelope: The nuclear envelope separates the contents of the nucleus from the cytoplasm, maintaining the nucleus as a distinct biochemical compartment houses the genetic material and serves as the site of transcription and RNA processing in eukaryotic cells 2
3 Figure 8.1 The nuclear envelope The inner nuclear membrane is lined by the nuclear lamina, which serves as an attachment site for chromatin Structure of the Nuclear Envelope: The nuclear envelope consists of the inner and outer nuclear membranes which are joined at nuclear pore complexes, and an underlying nuclear lamina Figure 8.2 Electron micrograph showing nuclear pores 3
4 Nuclear Lamina: a fibrous meshwork that provides structural support to the nucleus The Nuclear Pore Complex: Nuclear pore complexes are large structures provide the only routes through which molecules can travel between the nucleus and cytoplasm Figure 8.2 Electron micrograph showing nuclear pores 4
5 Figure 8.7 Model of the nuclear pore complex The complex consists of an assembly of 8 spokes attached to rings on the cytoplasmic & nuclear sides of the nuclear envelope Figure 8.5 Molecular traffic through nuclear pore complexes Small molecules are able to diffuse freely through open channels in the nuclear pore complex. Macromolecules are selectively transported in an energy-dependent process. 5
6 Selective Transport of Proteins to and from the Nucleus: Proteins destined for import to the nucleus contain nuclear localization signals that are recognized by receptors that direct transport through the nuclear pore complex. Figure 8.8 Nuclear localization signals Two types of nuclear transport receptor (karyopherins): importins & exportins 6
7 Selective Transport of Proteins to and from the Nucleus The small GTP-binding protein Ran is required for translocation through the nuclear pore complex and determines the directionality of transport. Ran GEF: guanine nucleotide exchange factor Ran GAP: GTPase-activating protein Figure 8.9 Protein import through the nuclear pore complex a = importin; b = Ran 7
8 Selective Transport of Proteins to and from the Nucleus Proteins that shuttle back and forth between the nucleus and the cytoplasm contain nuclear export signals that target them for transport from the nucleus to the cytoplasm. Figure 8.10 Nuclear export Complexes between cargo proteins bearing nuclear export singals (NES), exportins, & Ran/GTP form in the nucleus 8
9 Regulation of Nuclear Protein Import: The activity of some proteins, such as transcription factors, is controlled by regulation of their import to the nucleus. Transport of RNAs: RNAs are transported through the nuclear pore complex as ribonucleoprotein complexes. Messenger RNAs, ribosomal RNAs, and transfer RNAs are exported from the nucleus to function in protein synthesis. Small nuclear RNAs are initially transported from the nucleus to the cytoplasm, where they associate with proteins to form snrnps; then they return to the nucleus. 9
10 Figure 8.14 Transport of snrnas between nucleus and cytoplasm Internal Organization of the Nucleus Organizes the genetic material & localizes nuclear functions to specific sites or domains 10
11 Chromosomes and Higher- Order Chromatin Structure: The interphase nucleus contains transcriptionally inactive, highly condensed heterochromatin as well as decondensed euchromatin. Chromosomes and Higher- Order Chromatin Structure: Interphase chromosomes are organized within the nucleus and divided into large looped domains that function as independent units. 11
12 Figure 8.18 Organization of chromosomes in the mammalian nucleus Individual chromosomes occupy distinct territories within the nuclei Functional Domains within the Nucleus: Clustered sites of DNA replication Localization of splicing comonents 12
13 THE NUCLEOLUS The site of rrna transcription & processing, & of ribosome assembly Ribosomal RNA Genes and the Organization of the Nucleolus: The nucleolus is organized around the genes for ribosomal RNAs. It is the site of rrna transcription and processing, and of ribosome assembly. 13
14 Figure 8.22 Ribosomal RNA genes Each rrna gene is a single transcription unit containing the 18S, 5.8S, & 28S rrnas & transcribed spacers. The rrna genes are organized in tandem arrays, separated by nontranscribed spacer DNA. Figure 8.23 Nucleoli in amphibian oocytes The amplified rrna genes of Xenopus oocytes are clustered in multiple nucleoli 2000-fold ~1 million copies per cell ribosomes per oocyte 14
15 Transcription and Processing of rrna: The primary transcript of the rrna genes is 45S pre-rrna, which is processed to yield 18S, 5.8S, and 28S rrnas. Figure 8.25 Transcription of rrna Figure 8.26 Processing of pre-rrna ETS: external transcribed spacers; ITS: internal - 15
16 Figure 8.27 Structure of the nucleolus FC: fibrillar certer; DFC: dense fibrillar component; G: granular component FC: rrna transcription DFC: processing G: ribosome assembly Transcription and Processing of rrna Processing of prerrna is mediated by small nucleolar RNAs (snornas) Catalyze methylation Figure 8.27 Role of snornas in base modification of pre-rrna 16
17 Ribosome Assembly: Ribosomal subunits are assembled within the nucleolus from rrnas and ribosomal proteins. THE NUCLEUS DURING MITOSIS Vertebrate Mitosis 1994 Jeremy Pickett-Heaps [vert_mitosis.mov] Mitosis and Nondisjunction Conly Rieder [mitosis.mov] 17
18 Figure 8.29 The nucleus during mitosis Figure 8.30 Closed & open mitosis Some unicellular eukaryotes (yeasts) 18
19 Dissolution of the Nuclear Envelope: Entry into mitosis is signaled by activation of the Cdc2 protein kinase. Dissolution of the Nuclear Envelope In most cells, the nuclear envelope breaks down at the end of prophase. Depolymerization of the nuclear lamina results from phosphorylation of the lamins by Cdc2 and other protein kinases. 19
20 Figure 8.32 Breakdown of the nuclear membrane As the nuclear lamina dissociates, the nuclear membrane fragments into vesicles. The B-type lamins remain bound to these vesicles, while lamins A & C are released as free dimers Chromosome Condensation: Phosphorylation of histones H1 and H3 is correlated with the condensation of mitotic chromosomes, and H3 phosphorylation is required for proper chromosome condensation. A complex of proteins called condensin is activated by Cdc2 phosphorylation and functions in chromosome condensation by wrapping the DNA into a compact structure. 20
21 Figure 8.33 Chromosome condensation Electron micrograph showing the condensation of individual chromosomes during the prophase of mitosis Re-formation of the Interphase Nucleus: Inactivation of Cdc2 at the end of mitosis leads to re-decondensation. Nuclear proteins are then selectively imported through nuclear pore complexes. 21
22 Figure 8.34 Re-formation of the nuclear envelope 1st step: binding of membrane vesicles to chromosome Figure 8.34 Reformation of the nuclear envelope The vesicles then fuse The nuclear lamina reassembles The chromosomes decondense 22
23 Nucleus and Nucleolus 1994 Jeremy Pickett-Heaps [nucl.mov] Embryonic Chromosomes and Spindles 1995 William Theurkauf [embryo_chrom.mov] 23
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