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You are watching: What forms around the chromatids during mitosis

Cooper GM. The Cell: A molecule Approach. 2nd edition. Sunderland (MA): Sinauer Associates; 2000.

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M phase is the most dramatic duration of the cell cycle, entailing a significant reorganization of basically all cell components. During mitosis (nuclear division), the chromosomes condense, the nuclear envelope of most cells breaks down, the cytoskeleton reorganizes to kind the mitotic spindle, and also the chromosomes move to opposite poles. Chromosome distinction is then usually adhered to by cell department (cytokinesis). Although countless of these events have been debated in ahead chapters with respect come the structure and function of the nucleus and also cytoskeleton, they room reviewed here in the paper definition of a coordinated view of M phase and also the action of MPF.

Stages the Mitosis

Although numerous of the details that mitosis vary among different organisms, the fundamental processes that ensure the faithful segregation of sister chromatids space conserved in all eukaryotes. These straightforward events that mitosis include chromosome condensation, formation of the mitotic spindle, and also attachment of chromosomes come the spindle microtubules. Sister chromatids then different from each other and move come opposite poles of the spindle, complied with by the formation of daughter nuclei.

Mitosis is conventionally divided into 4 stages—prophase, metaphase, anaphase, and also telophase—which are depicted for an pet cell in numbers 14.23 and 14.24. The start of prophase is marked by the figure of condensed chromosomes, every of which consists of two sister chromatids (the daughter DNA molecules developed in S phase). These freshly replicated DNA molecule remain intertwined throughout S and also G2, coming to be untangled throughout the process of chromatin condensation. The condensed sister chromatids space then organized together in ~ the centromere, i m sorry (as debated in chapter 4) is a DNA succession to i m sorry proteins bind to type the kinetochore—the website of eventual attachment of the spindle microtubules. In enhancement to chromosome condensation, cytoplasmic transforms leading to the development of the mitotic spindle initiate throughout prophase. The centrosomes (which had duplicated during interphase) separate and also move to opposite sides of the nucleus. There they serve as the two poles the the mitotic spindle, which starts to kind during late prophase.


Figure 14.23

Stages the mitosis in an animal cell. Throughout prophase, the chromosomes condense and centrosomes relocate to opposite sides of the nucleus, initiating development of the mitotic spindle. Break down of the atom envelope then allows spindle microtubules to attach (more...)


Figure 14.24

Fluorescence micrographs the chromatin, keratin, and also microtubules during mitosis the newt lung cells. Chromatin is stained blue, keratin is stained red, and microtubules room stained green. (Conly L. Rieder/ biological Photo Service.)

In greater eukaryotes the finish of prophase synchronizes to the failure of the nuclear envelope. As discussed in chapter 8, however, nuclear envelope breakdown is not a universal feature of mitosis. In particular, yeasts and many various other unicellular eukaryotes undergo “closed mitosis,” in i m sorry the nuclear envelope remains intact (see figure 8.30). In these cells the spindle pole bodies are embedded within the nuclear envelope, and the cell nucleus divides in two following migration of daughter chromosomes come opposite poles that the spindle.

Following perfect of prophase, the cell enters prometaphase—a transition duration between prophase and metaphase. During prometaphase the microtubules the the mitotic spindle connect to the kinetochores the condensed chromosomes. The kinetochores the sister chromatids are oriented ~ above opposite political parties of the chromosome, so they attach to microtubules create from opposite poles the the spindle. The chromosomes shuffle ago and forth till they eventually align on the metaphase key in the facility of the spindle. In ~ this stage, the cell has reached metaphase.

Most cell remain just briefly in ~ metaphase before proceeding to anaphase. The change from metaphase come anaphase is triggered by breakage of the link between sister chromatids, which then separate and also move come opposite poles the the spindle. Mitosis ends v telophase, throughout which nuclei re-form and the chromosomes decondense. Cytokinesis usually begins during late anaphase and is virtually complete through the end of telophase, resulting in the development of 2 interphase daughter cells.

MPF and also Progression to Metaphase

Mitosis requires dramatic transforms in multiple to move components, resulting in a major reorganization of the entire structure that the cell. As disputed earlier in this chapter, these occasions are initiated by activation the the MPF protein kinase (Cdc2/cyclin B). It shows up that MPF not only acts as a grasp regulator of the M step transition, phosphorylating and also activating various other downstream protein kinases, but additionally acts straight by phosphorylating few of the structure proteins connected in this cellular reorganization (Figure 14.25).


Figure 14.25

Targets of MPF. MPF cause multiple nuclear and also cytoplasmic transforms at the start of M phase, both through activating other protein kinases and also by phosphorylating protein such together condensins and the nuclear lamins.

The condensation of interphase dyed to kind the compact chromosomes of mitotic cells is a vital event in mitosis, vital in enabling the chromosomes to move along the mitotic spindle without coming to be broken or tangled v one another. As debated in thing 4, the dyed in interphase nuclei condenses nearly a thousand fold during the formation of metaphase chromosomes. Such very condensed chromatin cannot be transcribed, so warrior ceases as chromatin condensation take away place. Despite the an essential importance the this event, we do not completely understand one of two people the structure of metaphase chromosomes or the molecular mechanism of chromatin condensation. However, protein complexes called condensins have recently been discovered to journey chromosome condensation by pack DNA roughly itself, compacting chromosomes right into the condensed mitotic structure. The condensins space phosphorylated straight by the Cdc2 protein kinase, i beg your pardon drives chromatin condensation through activating condensins together cells enter mitosis. One molecular change that usually accompanies chromosome condensation is phosphorylation of histone H1, so it is notable that histone H1 is likewise a substrate for Cdc2. However, histone H1 phosphorylation is not compelled for mitotic chromosome condensation, so the significance of H1 phosphorylation by Cdc2 is unclear. In contrast, chromosome condensation has been presented to call for phosphorylation of histone H3. Possibly surprisingly, however, histone H3 is no phosphorylated through Cdc2 and the kinase responsible because that H3 phosphorylation in mitotic cells stays to it is in identified.

Breakdown of the nuclear envelope, which is one of the many dramatic occasions of mitosis, to represent the most clearly defined target for MPF action. As questioned in chapter 8, Cdc2 phosphorylates the lamins, leading straight to depolymerization the the atom lamina (see number 8.31). This is followed by fragmentation of the nuclear membrane into tiny vesicles, which ultimately fuse to form new daughter nuclei at telophase. The endoplasmic reticulum and Golgi apparatus similarly fragment into small vesicles, which can then be distributed to daughter cells at cytokinesis. The failure of these membranes is likewise induced through MPF, and also may in part be mediated by Cdc2 phosphorylation that the Golgi matrix protein GM130, i m sorry is required for the docking of COPI-coated vesicles to the Golgi membrane. Phosphorylation and also inactivation the GM130 through Cdc2 inhibits vesicle docking and also fusion, causing fragmentation the the Golgi apparatus. However, extr targets the Cdc2 may additionally be involved, and also the mechanisms through which MPF leads to membrane fragmentation remain to be fully elucidated.

The reorganization that the cytoskeleton that culminates in development of the mitotic spindle outcomes from the dynamic instability that microtubules (see chapter 11). In ~ the beginning of prophase, the centrosomes move to opposite sides of the nucleus. The climb in MPF task then induces a dramatic adjust in the dynamic actions of microtubules. First, the rate of microtubule disassembly increases, leading to depolymerization and also shrinkage of the interphase microtubules. This disassembly is thought to an outcome from phosphorylation that microtubule-associated proteins, either by MPF chin or by other MPF-activated protein kinases. In addition, the number of microtubules create from the centrosomes increases, therefore the interphase microtubules are replaced by large numbers of brief microtubules radiating from the centrosomes.

The failure of the nuclear envelope then enables some the the spindle microtubules to connect to chromosomes at their kinetochores (Figure 14.26), initiating the process of chromosome activity that characterizes prometaphase. The proteins assembled in ~ the kinetochore include microtubule motors that straight the movement of chromosomes toward the minus end of the spindle microtubules, which room anchored in the centrosome. The activity of this proteins, which draw chromosomes towards the centrosome, is protest by the expansion of the spindle microtubules, i beg your pardon pushes the chromosomes away from the spindle poles. Consequently, the chromosomes in prometaphase shuffle back and forth between the centrosomes and also the facility of the spindle.


Figure 14.26

Electron micrograph of microtubules attached come the kinetochore the a chromosome. (Conly L. Rieder/ biological Photo Service.)

Microtubules indigenous opposite poles the the spindle eventually connect to the 2 kinetochores of sister chromatids (which are situated on opposite political parties of the chromosome), and also the balance of forces acting top top the chromosomes leader to their alignment on the metaphase plate in the facility of the spindle (Figure 14.27). As discussed in thing 11, the spindle is composed of both kinetochore microtubules, which space attached to the chromosomes, and polar microtubules, i m sorry overlap v one an additional in the center of the cell. In addition, short astral microtubules radiate external from the centrosomes towards the cell periphery.

Figure 14.27

The metaphase spindle. (A) The spindle consists of three kinds that microtubules. Kinetochore microtubules are attached come chromosomes, polar microtubules overlap in the center of the cell, and astral microtubules radiate from the centrosome to the cabinet (more...)

Proteolysis and also the Inactivation of MPF: Anaphase and also Telophase

As questioned earlier in this chapter, vital cell cycle checkpoint monitors the alignment the chromosomes top top the metaphase spindle. As soon as this has actually been accomplished, the cell proceeds to initiate anaphase and complete mitosis. The progression from metaphase to anaphase results from ubiquitin-mediated proteolysis of vital regulatory proteins, triggered by activation that a ubiquitin ligase (see number 7.39) called the anaphase-promoting complex. Activation that the anaphase-promoting complex is induced by MPF at the beginning of mitosis, so MPF eventually triggers its very own destruction. The anaphase-promoting facility remains inhibited, however, until the cell passes the metaphase checkpoint, after i beg your pardon activation of the ubiquitin deterioration system brings about the transition from metaphase come anaphase and progression with the rest of mitosis.

Activation that the anaphase-promoting complicated leads come the degradation of at the very least two key regulatory proteins (Figure 14.28). The onset of anaphase results from proteolytic destruction of a protein called Scc1, a ingredient of a facility of proteins called cohesins that maintain the connection between sister chromatids while they room aligned on the metaphase plate. Degradation of Scc1 is no catalyzed straight by the anaphase-promoting complex, which instead degrades a regulation protein called Pds1. Deterioration of Pds1 consequently activates another protein, referred to as Esp1, which leader to proteolysis of the cohesin Scc1. Cleavage of Scc1 breaks the linkage between sister chromatids, allowing them come segregate by relocating to opposite poles that the spindle (Figure 14.29). The separation that chromosomes during anaphase climate proceeds together a result of the activity of several species of engine proteins associated with the spindle microtubules (see numbers 11.48 and also 11.49).

Figure 14.28

Targets the the cyclin B proteolysis system. The anaphase-promoting complicated is a ubiquitin ligase the is activated following passage through the metaphase checkpoint. Its activation brings about the shift from metaphase to anaphase by leading to (more...)

The other vital regulatory protein targeted for ubiquitination and also degradation by the anaphase-promoting complicated is cyclin B. Deterioration of cyclin B leads to inactivation of MPF, i m sorry is required for the cabinet to leave mitosis and return to interphase. Countless of the moving changes connected in these transitions are simply the reversal the the occasions induced through MPF throughout entry right into mitosis. For example, reassembly that the atom envelope, dyed decondensation, and the return the microtubules to an interphase state probably an outcome directly native loss the MPF activity and dephosphorylation of protein that had actually been phosphorylated by MPF at the beginning of mitosis. As discussed next, inactivation the MPF additionally triggers cytokinesis.


The perfect of mitosis is normally accompanied by cytokinesis, providing rise to two daughter cells. Cytokinesis usually initiates in so late anaphase and also is motivated by the inactivation the MPF, in order to coordinating nuclear and cytoplasmic division of the cell. As questioned in chapter 11, cytokinesis of animal cells is mediated by a contractile ring that actin and also myosin II filaments that develops beneath the plasma membrane (Figure 14.30). The location of this ring is established by the position of the mitotic spindle, so the cabinet is at some point cleaved in a airplane that passes with the metaphase key perpendicular to the spindle. Cleavage proceeds as contraction the the actin-myosin filaments pulls the plasma membrane inward, at some point pinching the cabinet in half.

Figure 14.30

Cytokinesis of pet cells. (A) Cytokinesis outcomes from contraction of a ring the actin and myosin filaments, i m sorry pinches the cabinet in two. (B) Scanning electron micrograph that a frog egg experience cytokinesis. (B, David M. Phillips/Visuals Unlimited). (more...)

The device of cytokinesis is different for greater plant cells, which are surrounded through rigid cabinet walls. Quite than being pinched in half by a contractile ring, these cells divide by forming brand-new cell walls and also plasma membranes inside the cabinet (Figure 14.31). In early telophase, vesicles moving cell wall surface precursors from the Golgi apparatus associate with spindle microtubules and also accumulate in ~ the former site the the metaphase plate. These vesicles then fuse to form a large, membrane-enclosed, disclike structure, and their polysaccharide contents assemble to type the procession of a brand-new cell wall surface (called a cell plate). The cabinet plate broadens outward, perpendicular to the spindle, until it get the plasma membrane. The membrane surrounding the cell plate climate fuses through the parental plasma membrane, separating the cabinet in two.

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Figure 14.31

Cytokinesis in higher plants. Golgi vesicles carrying cell wall surface precursors associate with polar microtubules in ~ the previous site of the metaphase plate. Blend of these vesicles returns a membrane-enclosed, disclike framework (the early on cell plate) the (more...)

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