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At the same time,the amount of y-tubulin ring complexes in each centrosome increases greatly,increasing the ability of the centrosomes to nucleate new microtubules, a processcalled centrosomematuration.Multiple motor proteins drive the separation of centrosomes in early mitosis. In prophase, minus-end directed dlnein motor proteins at the plus ends ofastral microtubules provide the major force.
These motors are anchored at thecell cortex or on the nuclear envelope, and their movement toward the microtubule minus end pulls the centrosomes apart (see Figure 17-30). Followingnuclear envelope breakdown at the end of prophase, interactions between thecentrosomal microtubules and the cell cortex allow actin-myosin bundles in thecortex to pull the centrosomes further apart. Finally, kinesin-5 motors cross-linkthe overlapping, antiparallel ends of interpolar microtubules and push the polesapart (seeFigure 17-30).The balance of opposing forces generated by different types of motor proteins determines the final length of the spindle.
Dynein and kinesin-s motorsgenerally promote centrosome separation and increase spindle length. Kinesin14 proteins do the opposite: they are minus-end directed motors and interactwith a microtubule from one pole while traveling toward the minus end of anantiparallel microtubule from the other pole; as a result, they tend to pull thepoles together. It is not clear how the cell regulates the balance of opposingforces to generate the appropriate spindle length (Figure 17-32).M-Cdk and other mitotic protein kinases are required for centrosome separation and maturation. M-Cdk and aurora-A phosphorylate kinesin-S motorsand stimulate them to drive centrosome separation.Aurora-A and PIk also phosphorylate components of the centrosome and thereby promote its maturation.TheCompletionof SpindleAssemblyin AnimalCellsRequiresNuclearEnvelopeBreakdownThe centrosomes and microtubules of animal cells are located in the cytoplasm,separated from the chromosomes by the double membrane barrier of thenuclear envelope (discussed in Chapter 12).
Clearly, the attachment of sisterchromatids to the spindle requires the removal of this barrier. In addition, manyof the motor proteins and microtubule regulators that promote spindle assembly are associatedwith the chromosomes inside the nucleus. Nuclear envelopebreakdor.rmallows these proteins to carry out their important functions in spindle assemblv.1080Chapter17:TheCell CycleNuclear envelope breakdown is a complex, multi-step process that isthought to begin when M-cdk phosphorylates several subunits of the giantnuclear pore complexes in the nuclear envelope. This initiates the disassemblyof nuclear pore complexes and their dissociation from the envelope.
M-Cdkalso phosphorylates components of the nuclear lamina, the structural framework that lies beneath the envelope. The phosphorylation of these lamina components and of several inner nuclear envelope proteins leads to disassembly ofthe nuclear lamina and the breakdown of the envelope membranes into smallvesicles.MicrotubuleInstabilitylncreasesGreatlyin MitosisMost animal cells in interphase contain a cytoplasmic array of microtubulesradiating out from the single centrosome.As discussedin chapter 16, the microtubules of this interphase array are in a state of dynamic instability, in whichindividual microtubules are either growing or shrinking and stochasticallyswitch between the two states.
The switch from growth to shrinkage is called acatastrophe, and the switch from shrinkage to growth is called a rescue(see Figure l6-16). New microtubules are continually being created to balance the lossof those that disappear completely by depolymerization.Entry into mitosis signals an abrupt change in the cell's microtubules. Theinterphase array of few, long microtubules is converted to a larger number ofshorter and more dynamic microtubules surrounding each centrosome. Duringprophase, and particularly in prometaphase and metaphase (seepanel l7-l), thehalf-life of microtubules decreases dramatically.
This increase in microtubuleinstability, coupled with the increased ability of centrosomes to nucleate microtubules as mentioned earlier, results in remarkably dense and dlmamic arrays ofspindle microtubules that are ideally suited for capturing sister chromatids.M-Cdk initiates these changes in microtubule behavior, at least in part, byphosphorylating two classesof proteins that control microtubule dynamics (discussed in chapter 16).
These include microtubule-dependent motor proteinsand microtubule-associated proteins (MAPs). Experiments using cell-freeXenopus egg extracts,which reproduce many of the changes that occur in intactcells during the cell cycle, have revealed the roles of these regulators in controlling microtubule dyrramics.If centrosomes and fluorescent tubulin are added tothese extracts,fluorescent microtubules nucleate from the centrosomes,and wecan observe the behavior of individual microtubules by time-lapse fluorescencevideo microscopy. The microtubules in mitotic extracts differ from those ininterphase extracts primarily by the increased rate of catastrophes,in which themicrotubules switch abruptly from slow growth to rapid shortening.TWo classes of proteins govern microtubule dlrramics in mitosis.
proteinscalled catastrophe factors destabilize microtubule arrays by increasing the frequency of catastrophes (see Figure 16-16). one of these proteins is a kinesinrelated protein that does not function as a motor. MAps, by contrast, have theopposite effect, stabilizing microtubules in various ways: they can increase the frequency of rescues,in which microtubules switch from shrinkage to growth, or theycan either increase the growth rate or decreasethe shrinkage rate of microtubules.Thus, in principle, changes in catastrophe factors and MAps can make microtubules much more dFramic in M phase by increasing total microtubule depolymerization rates, decreasing total microtubule polymerization rates, or both.constant throughout the cell cycle, the balance between the two opposing activities of the MAP and catastrophe factor would shift, increasing the dynamicinstability of the microtubules.'seluosoJlual ululuoc ,{lerurou }Eql sllaJeql azrue8Jodlaq saurosouloJqJ aqlur ua1g '(gt-ZI arnglg) saluosoJlue3lnoqllv\ dola,rap so^rqlua clueSouaqUgdaseq] ur selpurds JItolIu eqt '(IZ Jalduq3 uI passnJslp) 33a ue sezITIUeJtl ueq^ errrosoJlual eql sapl^oJdri.lprurou ulJeds etqlsc-:(/ttlDJ!rcuaSouaquad'srteqt) 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