Page "Spindle checkpoint" Paragraph 33
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The leading model of MCC formation is the " MAD2-template model ", which depends on the kinetochore dynamics of MAD2 to create the MCC.
Cells lacking Aurora B fail to arrest in metaphase even when chromosomes lack microtubule attachment.
Unattached kinetochores first bind to a MAD1-C-MAD2-p31 < sup > comet </ sup > complex and releases the p31 < sup > comet </ sup > through unknown mechanisms.
The resulting MAD-C-MAD2 complex recruits the open conformer of Mad2 ( O-Mad2 ) to the kinetochores.
This Mad1 / C-Mad2 complex is responsible for the recruitment of more O-Mad2 to the kinetochores, which changes its conformation to C-Mad2 and binds Cdc20 in an auto-amplification reaction.
Since MAD1 and CDC20 both contain a similar MAD2-binding motif, the empty O-MAD2 conformation changes to C-MAD2 while binding to CDC20.
This positive feedback loop is negatively regulated by p31 < sup > comet </ sup >, which competitively binds to C-MAD2 bound to either MAD1 or CDC20 and reduces further O-MAD2 binding to C-MAD2.
Further control mechanisms may also exist, considering that p31 < sup > comet </ sup > is not present in lower eukaryotes.
The ' template model ' nomenclature is thus derived from the process where MAD1-C-MAD2 acts as a template for the formation of C-MAD2-CDC20 copies.
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