7.10
During mitosis, the bioriented chromosomes oscillate to stabilize their spindle attachments and subsequently assemble along the metaphase plate at the equator of the mitotic spindle.
Multiple forces act on chromosomes after they are attached to the mitotic spindle.
A major poleward force exerted along the kinetochore-microtubule, as a result of plus-end de-polymerization, pulls the kinetochore and its associated chromosome towards the spindle pole. At the kinetochore, Ndc80 protein complexes link the kinetochore to the microtubule through multiple low-affinity attachments along microtubule sides.
During microtubule plus-end de-polymerization, the Ndc80 attachments break and reform at new sites to maintain the kinetochore-microtubule connection. The mechanism gradually pulls the chromosome towards the spindle pole, as the microtubule shortens in length.
A second poleward force results from microtubule flux. Microtubule minus-end depolymerization generates a minus-end directed flux, causing microtubule movement towards the spindle pole. Coordinating plus-end polymerization compensates for the minus-end depolymerization, allowing microtubules to maintain their length.
A third force, the polar ejection force or polar wind, generated by kinesin-4 and 10 motor proteins pushes the chromosomes away from the spindle poles. Kinesin-4 and 10 link chromosomal arms with interpolar microtubules. These plus-end directed motor proteins move the chromosome toward the spindle equator.
A balanced interplay of these multiple opposing forces enables the bioriented chromosomes to precisely align along the metaphase plate, in preparation for chromosomal segregation.
During mitosis, chromosome movements occur through the interplay of multiple piconewton level forces. In prometaphase, these forces help in chromosome…
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