Microtubule turnover as a mechanism of mitosis and its possible evolution

Movement of chromosomes during mitosis appears to be coupled to the unidirectional turnover of spindle microtubules. This paper outlines a directional turnover model of mitosis which hypothesizes that: (1) Unidirectional turnover of tubulin subunits and microtubule-associated proteins occurs from an assembly site at one end to a disassembly site at the other end of mitotic apparatus microtubules. (2) The components of interpolar microtubules are continuously moving toward each pole due to their assembly in the opposite half spindle and disassembly in the near half spindle. (3) Mitotic chromosome movements are coupled to this directional turnover by selective formation of semi-stable lateral interactions or bridges between kinetochore microtubules and parallel interpolar microtubules. (4) The anaphase velocity of kinetochores relative to the poles is determined by the rate that kinetochore microtubules disassemble on their poleward ends minus the rate they assemble at their kinetochore ends. (5) Spindle elongation occurs when assembly of interpolar microtubules is more rapid than their disassembly or when interpolar microtubules slide in an anti-parallel arrangement. (6) The velocity of chromosome separation is the sum of spindle elongation and the poleward movements of sister chromosomes. Evidence for and against these hypotheses and some possible steps in the evolution of this type of mechanism are discussed.