Ultrastructural studies in the Rhodophyta. I. Development of mitotic spindle poles in Apoglossum ruscifolium, Kylin

Abstract

The ultrastructure of somatic mitosis in the germlings of Apoglossum ruscifolium, Kylin, fixed from its natural habitat, was examined to investigate spindle-pole development and the role of the ‘polar ring’. It was found that the polar ring originates from a special nuclear pore raised above a small projection of the nuclear envelope. The initiation of mitotic polarity takes the form of changes in the nuclear envelope. These are: close crowding of pores, heavy deposition of electron-opaque material and attachment of microtubules. No such phenomena are to be observed in the equatorial regions of the nuclear envelope, which persists throughout mitosis. The next stage is the development of ‘clear zones’, which become filled with microtubules excluding all other structures, notably ribosomes, at both poles. At pre-metaphase, microtubules begin to be extended from the clear zones through polar fenestrae in the nuclear envelope into the nucleus itself. With subsequent development of the intranuclear spindle, the microtubules in the clear zones show signs of degeneration. At metaphase, the polar regions of the nuclear envelope begin to return to their normal condition. After metaphase, the polar rings degenerate. Thus it is primarily the nuclear envelope, via its polar modifications, which begins the organization of the mitotic spindle. The capabilities of nuclear pores are summarized. In the Discussion, the polar rings are compared with procentrioles; since they appear to have a passive role in spindle-pole organization, it is suggested that they may represent an evolutionary stage prior to the procentriole. The origin of the centriole ultimately from a nuclear pore is presented as an hypothesis. It is concluded to be unlikely that the polar ring is a degenerate centriole, and therefore it is proposed that the red algae never had centrioles in the course of their evolution. Thus the view of the non-flagellate ancestry of the red algae is supported.