Evidence for polewards forces on chromosome arms during anaphase

We have identified new mitotic forces in crane-fly spermatocytes, separate from forces on the kinetochore, that propel chromosome arms in anaphase towards the spindle pole. In normal spermatocytes, the chromosome arms in anaphase generally trail the kinetochore to the pole. After ultraviolet-microbeam irradiation of a kinetochore spindle fibre, however, chromosome arms moved closer to the pole than the kinetochore. This poleward arm-movement occurred regardless of whether the irradiation stopped the movement of the associated chromosomes, and occurred both in chromosomes associated with the irradiated fibre and in chromosomes not associated with the irradiated fibre. Arms that moved ahead of the kinetochore continued to lead the kinetochore to the pole for the duration of anaphase. Ultraviolet-microbeam-irradiation-induced movement of arms ahead of the kinetochore is specific for irradiation of spindle fibres: irradiations of the cytoplasm outside the spindle had no effect, and irradiations of the region between spindle and mitochondrial sheath (that outlines the spindle) and irradiations of the interzonal region are much less effective than irradiations of spindle fibres in causing arms to move. We argue that in crane-fly spermatocytes forces propelling chromosome arms toward the pole are part of normal anaphase.