Meiosis in Sciara coprophila: structure of the spindle and chromosome behavior during the first meiotic division.

Light microscope descriptions of meiosis I in males of the fungus gnat Sciara coprophila suggested the presence of a monopolar spindle in which maternal and limited chromosomes move poleward while paternal chromosomes "back away" from the pole. The ultrastructural analysis reported here, based upon serial sections of cells in different stages of meiosis I, shows that the spindle is indeed monopolar with a distinctive differentiation, the polar complex, at one pole. This complex is the focus of a conical radiation of spindle microtubules. Kinetochores of paternal chromosomes face the complex and microtubules associated with these kinetochores run toward the complex. No kinetochore microtubules were discovered on maternal or limited chromosomes. When the position of paternal, maternal, and limited chromosomes is compared at various stages, it is found that limited chromosomes always remain near the polar complex, paternal chromosomes remain far from it and only maternal chromosomes move closer to the pole. Apparently, chromosome segregation does not depend on paternal chromosomes "backing away" from the pole, and the required movement of maternal chromosomes take place in the absence of kinetochore microtubules. In the prophase nucleus, limited and maternal chromosomes are already spatially separate from paternal chromosomes before the spindle forms. Thus, the monopolar spindle functions only to increase the distance between already segregated sets of chromosomes. An extensive system of microtubule-associated membranes outlines the spindle; the possibility that maternal chromosome movement is somehow related to the presence of this membrane is discussed.