Cell Division in Oedogonium II. Nuclear Division in O. Cardiacum

Abstract

Nuclear division in O. cardiacum is described. Before division, the nucleus enlarges considerably. At prophase, the nucleolus starts dispersing and kinetochores appear on the condensing chromatin, situated and oriented apparently at random in the nucleus. By prometaphase, the kinetochore pairs become aligned along the spindle axis before moving into the metaphase-plate configuration; this supports an earlier theory explaining metakinesis. During prophase and metaphase particularly, the nuclear envelope at the poles forms channels that extend for some distance into the cytoplasm; these may also bifurcate. The nucleolus disperses but remains in the intranuclear spindle throughout division as a loosely knit skein of granular material. The kinetochores have a complex structure, up to seven distinct layers being detectable; the kinetochore pairs split, and then migrate polewards at anaphase with the rest of the chromosome trailing behind. Large numbers of microtubules run from the kinetochore into evaginations of the nuclear envelope which increase in size during anaphase. The spindle grows in length considerably during anaphase, this coinciding with a proliferation of interzonal microtubules, first seen amongst the trailing chromosome arms. The nuclear envelope enclosing the spindle becomes severely stretched at this stage; it contracts closely around each of the daughter nuclei, isolating them from the rest of the spindle (including microtubules and the remains of the nucleolus). The spindle then collapses; the nuclei come together and then flatten against one another; between them, vesicles and other components of the septum collect amongst a large number of transversely oriented micro tubules.