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You are watching: During which phase of mitosis do the nuclear envelope and nucleoli disappear?

Cooper GM. The Cell: A molecule Approach. Second edition. Sunderland (MA): Sinauer Associates; 2000.


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A distinct feature the the nucleus is that it disassembles and re-forms every time most cells divide. In ~ the beginning of mitosis, the chromosomes condense, the core source disappears, and the nuclear envelope breaks down, leading to the release of many of the components of the nucleus into the cytoplasm. In ~ the end of mitosis, the process is reversed: The chromosomes decondense, and also nuclear envelopes re-form roughly the separated set of daughter chromosomes. Chapter 14 gift a an extensive discussion the mitosis; in this ar we will consider the mechanisms affiliated in the disassembly and also re-formation that the nucleus. The procedure is managed largely through reversible phosphorylation and dephosphorylation of nuclear proteins result from the activity of the Cdc2 protein kinase, which is a critical regulator that mitosis in all eukaryotic bio cells.


Dissolution of the nuclear Envelope

In many cells, the disassembly of the nuclear envelope marks the end of the prophase the mitosis (Figure 8.29). However, this disassembly that the cell core is not a universal function of mitosis and also does not take place in every cells. Part unicellular standard scale (e.g., yeasts) experience so-called close up door mitosis, in i m sorry the nuclear envelope remains undamaged (Figure 8.30). In close up door mitosis, the daughter chromosomes move to the contrary poles of the nucleus, which climate divides in two. The cell of greater eukaryotes, however, typically undergo open mitosis, which is characterized by failure of the atom envelope. The daughter chromosomes then move to the contrary poles the the mitotic spindle, and new nuclei reassemble about them.


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Figure 8.29

The nucleus throughout mitosis. Micrographs illustrating the steady stages the mitosis in a plant cell. During prophase, the chromosomes condense, the nucleolus disappears, and the nuclear envelope division down. In ~ metaphase, the condensed chromosomes (more...)


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Figure 8.30

Closed and open mitosis. In closed mitosis, the atom envelope stays intact and also chromosomes move to opposite poles the a spindle within the nucleus. In open up mitosis, the atom envelope breaks down and also then re-forms approximately the two sets of be separate (more...)


Disassembly that the atom envelope, which parallels a similar breakdown of the endoplasmic reticulum, involves changes in all 3 of that is components: The nuclear membranes are fragmented into vesicles, the atom pore complexes dissociate, and the atom lamina depolymerizes. The finest understood of these events is depolymerization that the nuclear lamina—the meshwork the filaments basic the nuclear membrane. The nuclear lamina is created of fibrous proteins, lamins, i beg your pardon associate v each other to kind filaments. Disassembly the the nuclear lamina outcomes from phosphorylation that the lamins, which causes the filaments to malfunction into separation, personal, instance lamin dimers (Figure 8.31). Phosphorylation that the lamins is catalyzed by the Cdc2 protein kinase, which was introduced in thing 7 (see number 7.40) and also will be discussed in information in chapter 14 together a main regulator that mitosis. Cdc2 (as well as other protein kinases triggered in mitotic cells) phosphorylates all the different species of lamins, and also treatment of secluded nuclei v Cdc2 has been presented to be sufficient to induce depolymerization of the atom lamina. Moreover, the requirement for lamin phosphorylation in the malfunction of the nuclear lamina has actually been demonstrated straight by the construction of mutant lamins that deserve to no longer be phosphorylated. As soon as genes encoding these mutant lamins were introduced right into cells, their expression was found to block normal failure of the nuclear lamina together the cells entered mitosis.


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Figure 8.31

Dissolution that the nuclear lamina. The atom lamina is composed of a meshwork the lamin filaments. At mitosis, Cdc2 and other protein kinases phosphorylate the lamins, resulting in the filaments to dissociate into cost-free lamin dimers.


In concert through dissolution the the nuclear lamina, the atom membrane pieces into motor (Figure 8.32). The B-type lamins remain connected with this vesicles, but lamins A and C dissociate from the atom membrane and also are exit as free dimers in the cytosol. This distinction arises since the B-type lamins are permanently modified by the enhancement of lipid (prenyl groups), conversely, the C-terminal prenyl teams of A- and also C-type lamins are gotten rid of by proteolysis adhering to their incorporation right into the lamina. The nuclear pore complexes additionally dissociate right into subunits together a an outcome of phosphorylation of numerous nuclear sharp proteins. Integral atom membrane protein are also phosphorylated at mitosis, and phosphorylation of this proteins may be necessary in vesicle formation as well as in dissociation that the nuclear membrane native both chromosomes and also the nuclear lamina.


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Figure 8.32

Breakdown that the atom membrane. As the atom lamina dissociates, the atom membrane pieces into vesicles. The B-type lamins continue to be bound to this vesicles, while lamins A and also C are released as complimentary dimers.


Chromosome Condensation

The other major change in atom structure during mitosis is chromosome condensation. The interphase chromatin, i m sorry is currently packaged right into nucleosomes, condenses approximately a thousandfold more to kind the compact chromosomes viewed in mitotic cells (Figure 8.33). This condensation is required to enable the chromosomes to relocate along the mitotic spindle without ending up being tangled or damaged during their circulation to daughter cells. DNA in this highly condensed state can no much longer be transcribed, so every RNA synthesis stops throughout mitosis. Together the chromosomes condense and also transcription ceases, the nucleolus also disappears.


Figure 8.33

Chromosome condensation. Electron micrograph showing the condensation of individual chromosomes during the prophase the mitosis. (K. G. Murti/Visuals Unlimited.)


The condensed DNA in metaphase chromosomes appears to be organized into big loops, each encompassing about a hundreds kilobases the DNA, which room attached come a protein frame (see number 4.13). Despite its basic importance, the system of chromosome condensation during mitosis is no understood. The an easy unit the chromatin framework is the nucleosome, which is composed of 146 base pairs that DNA wrapped about a histone main point containing 2 molecules every of histones H2A, H2B, H3, and H4 (see figure 4.8). One molecule the histone H1 is bound to the DNA as it enters every nucleosome main point particle, and also interactions in between these H1 molecules are associated in the urgent of chromatin into higher-order, an ext compact structures. Histone H1 is a substrate for the Cdc2 protein kinase and is phosphorylated during mitosis of most cells, continual with the phosphorylation playing a duty in mitotic chromosome condensation. However, recent experiments have presented that phosphorylation the histone H1 is not compelled for chromosome condensation, so the potential duty of H1 phosphorylation is unclear. In contrast, phosphorylation that histone H3 has actually been found to be required for condensation of mitotic chromosomes, back the mechanism whereby H3 phosphorylation affect chromosome condensation stays to it is in elucidated.

Recent studies have additionally identified protein complexes referred to as condensins the play a major role in chromosome condensation. Condensins are forced for chromosome condensation in extracts of mitotic cells and appear to duty by wrapping DNA about itself, thereby compacting chromosomes right into the condensed mitotic structure. Condensins are phosphorylated and activated by the Cdc2 protein kinase, giving a direct link in between activation that Cdc2 and mitotic chromosome condensation.


Re-formation of the Interphase Nucleus

During the perfect of mitosis (telophase), two new nuclei kind around the separated set of daughter chromosomes (see number 8.29). Chromosome decondensation and also reassembly that the nuclear envelope show up to be signaled through inactivation that Cdc2, which was responsible because that initiating mitosis through phosphorylating to move target proteins, including the lamins, histone H3, and condensins. The progression from metaphase come anaphase involves the activation the a ubiquitin-mediated proteolysis system that inactivates Cdc2 by degrading its regulation subunit, cyclin B (see figure 7.40). Inactivation of Cdc2 leads to the dephosphorylation the the proteins that were phosphorylated at the initiation the mitosis, result in leave from mitosis and also the re-formation of interphase nuclei.

The initial step in re-formation of the atom envelope is the binding the the motor formed throughout nuclear membrane malfunction to the surface of chromosomes (Figure 8.34). This communication of membrane vesicles through chromosomes might be mediated by both lamins and integral membrane protein of the inner atom membrane. The vesicles then fuse to type a double membrane roughly the chromosomes. This is followed by reassembly of the atom pore complexes, re-formation the the atom lamina, and also chromosome decondensation. The vesicles an initial fuse to type membranes approximately individual chromosomes, which then fuse through each various other to kind a complete solitary nucleus.


Figure 8.34

Re-formation that the atom envelope. The very first step in reassembly that the atom envelope is the binding of membrane vesicles come chromosomes, which might be mediated by both integral membrane proteins and also B-type lamins. The vesicles then fuse, the nuclear (more...)


The initial re-formation the the nuclear envelope approximately condensed chromosomes excludes cytoplasmic molecules from the freshly assembled nucleus. The brand-new nucleus is climate able to increase via the selective income of atom proteins indigenous the cytoplasm. Because nuclear localization signals are not cleaved from proteins that space imported to the nucleus, the very same nuclear proteins the were released right into the cytoplasm following disassembly that the nuclear envelope in ~ the beginning of mitosis deserve to be reimported right into the new nuclei developed after mitosis. The nucleolus, too, re-forms together the chromosomes decondense and transcription of the rRNA gene begins, completing the return native mitosis to an interphase nucleus.

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