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You are watching: During which stage of mitosis do nucleoli reappear

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


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A distinct feature the the nucleus is that it disassembles and also re-forms each time many cells divide. At the beginning of mitosis, the chromosomes condense, the nucleolus disappears, and the atom envelope division down, bring about the release of most of the materials of the nucleus into the cytoplasm. At the finish of mitosis, the process is reversed: The chromosomes decondense, and also nuclear envelopes re-form about the separated sets of daughter chromosomes. Chapter 14 presents a comprehensive discussion the mitosis; in this ar we will consider the mechanisms connected in the disassembly and re-formation that the nucleus. The process is regulated largely by reversible phosphorylation and dephosphorylation of nuclear proteins resulting from the action of the Cdc2 protein kinase, i beg your pardon is a an essential regulator of mitosis in all eukaryotic bio cells.


Dissolution of the atom Envelope

In many cells, the disassembly that the nuclear envelope clues the end of the prophase that mitosis (Figure 8.29). However, this disassembly the the cell core is not a universal attribute of mitosis and does not take place in every cells. Some unicellular eukaryotes (e.g., yeasts) undergo so-called closed mitosis, in which the nuclear envelope remains undamaged (Figure 8.30). In close up door mitosis, the daughter chromosomes migrate to opposite poles of the nucleus, which climate divides in two. The cells of greater eukaryotes, however, normally undergo open mitosis, which is identified by failure of the atom envelope. The daughter chromosomes then migrate to opposite poles of the mitotic spindle, and brand-new nuclei reassemble approximately them.


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Figure 8.29

The nucleus throughout mitosis. Micrographs showing the gradual stages of mitosis in a plant cell. Throughout prophase, the chromosomes condense, the nuclear source disappears, and the nuclear envelope division down. At metaphase, the condensed chromosomes (more...)


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Figure 8.30

Closed and open mitosis. In closeup of the door mitosis, the atom envelope remains intact and also chromosomes migrate to opposite poles the a spindle within the nucleus. In open up mitosis, the atom envelope breaks down and then re-forms around the 2 sets of be separate (more...)


Disassembly of the atom envelope, i m sorry parallels a similar malfunction of the absorbent reticulum, involves transforms in all three of that components: The atom membranes are fragmented into vesicles, the atom pore complexes dissociate, and the atom lamina depolymerizes. The finest understood that these occasions is depolymerization that the atom lamina—the meshwork the filaments basic the atom membrane. The atom lamina is written of fibrous proteins, lamins, which associate through each other to kind filaments. Disassembly that the atom lamina outcomes from phosphorylation of the lamins, which reasons the filaments to malfunction into separation, personal, instance lamin dimers (Figure 8.31). Phosphorylation the the lamins is catalytic analysis by the Cdc2 protein kinase, which was presented in thing 7 (see figure 7.40) and will be discussed in detail in chapter 14 together a central regulator of mitosis. Cdc2 (as fine as various other protein kinases triggered in mitotic cells) phosphorylates every the different species of lamins, and treatment of diverted nuclei with Cdc2 has actually been presented to be enough to induce depolymerization that the atom lamina. Moreover, the need for lamin phosphorylation in the malfunction of the nuclear lamina has been demonstrated directly by the building of mutant lamins that deserve to no much longer be phosphorylated. Once genes encoding these mutant lamins to be introduced into cells, their expression was uncovered to block normal failure of the nuclear lamina as the cells entered mitosis.


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Figure 8.31

Dissolution the the nuclear lamina. The atom lamina consists of a meshwork of lamin filaments. At mitosis, Cdc2 and other protein kinases phosphorylate the lamins, leading to the filaments come dissociate into free lamin dimers.


In concert v dissolution the the nuclear lamina, the atom membrane fragments into vesicles (Figure 8.32). The B-type lamins remain connected with these vesicles, yet lamins A and also C dissociate native the nuclear membrane and are released as cost-free dimers in the cytosol. This difference arises since the B-type lamins are permanently modification by the addition of lipid (prenyl groups), vice versa, the C-terminal prenyl teams of A- and C-type lamins are eliminated by proteolysis complying with their incorporation right into the lamina. The atom pore complexes additionally dissociate right into subunits as a an outcome of phosphorylation of number of nuclear spicy proteins. Integral atom membrane protein are also phosphorylated in ~ mitosis, and phosphorylation of these proteins may be necessary in vesicle formation and in dissociation that the atom membrane native both chromosomes and also the nuclear lamina.


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Figure 8.32

Breakdown the the atom membrane. Together the atom lamina dissociates, the atom membrane fragments into vesicles. The B-type lamins continue to be bound to these vesicles, if lamins A and also C are released as cost-free dimers.


Chromosome Condensation

The other significant change in nuclear structure throughout mitosis is chromosome condensation. The interphase chromatin, i m sorry is already packaged right into nucleosomes, condenses roughly a thousandfold more to form the compact chromosomes checked out in mitotic cells (Figure 8.33). This condensation is needed to enable the chromosomes to relocate along the mitotic spindle without becoming tangled or damaged during their circulation to daughter cells. DNA in this extremely condensed state deserve to no longer be transcribed, so all RNA synthetic stops during mitosis. Together the chromosomes condense and transcription ceases, the nucleolus also disappears.


Figure 8.33

Chromosome condensation. Electron micrograph mirroring the condensation of individual chromosomes throughout the prophase that mitosis. (K. G. Murti/Visuals Unlimited.)


The condensed DNA in metaphase chromosomes shows up to be organized into large loops, each encompassing about a hundreds kilobases that DNA, which room attached to a protein scaffold (see number 4.13). In spite of its fundamental importance, the device of chromosome condensation throughout mitosis is no understood. The simple unit of chromatin framework is the nucleosome, which is composed of 146 basic pairs that DNA wrapped about a histone main point containing 2 molecules every of histones H2A, H2B, H3, and also H4 (see figure 4.8). One molecule that histone H1 is bound to the DNA as it enters each nucleosome core particle, and also interactions in between these H1 molecule are connected in the urgently of chromatin right into higher-order, an ext compact structures. Histone H1 is a substrate for the Cdc2 protein kinase and also is phosphorylated during mitosis of many cells, continual with the phosphorylation playing a role in mitotic chromosome condensation. However, current experiments have shown that phosphorylation the histone H1 is not required for chromosome condensation, for this reason the potential duty of H1 phosphorylation is unclear. In contrast, phosphorylation of histone H3 has been found to be forced for condensation that mitotic chromosomes, return the mechanism whereby H3 phosphorylation affects chromosome condensation remains to it is in elucidated.

Recent researches have likewise identified protein complexes dubbed condensins the play a major role in chromosome condensation. Condensins are compelled for chromosome condensation in extracts that mitotic cells and also appear to role by pack DNA roughly itself, in order to compacting chromosomes into the condensed mitotic structure. Condensins room phosphorylated and activated by the Cdc2 protein kinase, offering a straight link in between activation that Cdc2 and mitotic chromosome condensation.


Re-formation of the Interphase Nucleus

During the perfect of mitosis (telophase), two brand-new nuclei form around the separated to adjust of daughter chromosomes (see figure 8.29). Chromosome decondensation and reassembly that the atom envelope show up to it is in signaled by inactivation that Cdc2, which to be responsible because that initiating mitosis by phosphorylating to move target proteins, consisting of the lamins, histone H3, and condensins. The development from metaphase to anaphase entails the activation the a ubiquitin-mediated proteolysis mechanism that inactivates Cdc2 by degrading its regulatory subunit, cyclin B (see figure 7.40). Inactivation that Cdc2 leads to the dephosphorylation of the proteins the were phosphorylated at the initiation of mitosis, result in departure from mitosis and the re-formation that interphase nuclei.

The initial step in re-formation the the nuclear envelope is the binding of the vesicles formed during nuclear membrane break down to the surface of chromosomes (Figure 8.34). This communication of membrane vesicles v chromosomes might be mediated by both lamins and also integral membrane protein of the inner atom membrane. The vesicles climate fuse to kind a dual membrane roughly the chromosomes. This is followed by reassembly the the atom pore complexes, re-formation of the atom lamina, and chromosome decondensation. The vesicles very first fuse to kind membranes approximately individual chromosomes, which then fuse through each other to type a complete single nucleus.


Figure 8.34

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


The early stage re-formation of the atom envelope roughly condensed chromosomes excludes cytoplasmic molecule from the newly assembled nucleus. The brand-new nucleus is climate able to broaden via the selective income of atom proteins from the cytoplasm. Since nuclear localization signals room not cleaved from protein that space imported come the nucleus, the same nuclear proteins that were released into the cytoplasm complying with disassembly of the nuclear envelope in ~ the beginning of mitosis deserve to be reimported right into the new nuclei formed after mitosis. The nucleolus, too, re-forms as the chromosomes decondense and also transcription of the rRNA genes begins, completing the return native mitosis to an interphase nucleus.

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