CELL DIVISION IN COSMARIUM BOTRYTIS1

Cell division in Cosmarium is described. Premitotic cells are very dense; the semicells, previously appressed to one another, separate slightly during entry into prophase. This separation coincides with deposition of a girdle of new wall material around the isthmus, where the 2 semicells are joined. Micro-tubules, abundant around the isthmus wall during interphase, all disappear during prophase; meanwhile, other microtubules proliferate outside the nuclear envelope. By metaphase, the nucleolus has dispersed, although remnants of it persist. The nuclear envelope breaks up, but some membranes coat metaphase chromosomes. The spindle, while being typical, is somewhat multipolar; microtubules, usually associated with elements of endoplasmic reticulum, are oriented toward numerous regions in the poles. During anaphase and telophase, these spindle tubules become increasingly directed toward a few discrete foci, and they persist after telophase. Meanwhile, the septum grows from the girdle of wall material to bisect the cell. After cytokinesis, some microtubules reappear near the isthmus, but only adjacent to the older, non-expanding semicell wall. Cell expansion then takes place, during which the nucleus, ensheathed in a complex microtubular system, moves into the forming semicell. Later, the chloroplast follows the nucleus and its 2 pyrenoids elongate and divide. When semicell expansion is complete, the chloroplast cleaves adjacent to the isthmus. The nucleus, now apparently not associated with microtubules, concurrently moves back into the isthmus. Continuous deposition of primary wall material accompanies cell expansion. Wall materials are apparently secreted as aggregates (perhaps derived from the contents of vesicles) adjacent to the plasmalemma, whose fibrous components become increasingly oriented in the outer layers of the wall by stretching. Late in semicell formation, this deposition ceases and during further expansion, the semi-cell develops a pattern of warts and ridges. Secondary wall deposition under the primary wall then follows, matching this pattern of ornamentation. In addition, numerous plugs of amorphous material capped by specialized regions of the plasmalemma, traverse the entire thickness of the secondary wall which becomes further thickened at these particular sites. The amorphous plugs presumably are eroded away later to form the mucilage pores of the vegetative cells. The wall microtubules gradually become more symmetrically arrayed around the isthmus as this new secondary wall thickens. These observations are discussed in comparison with other work on morphogenesis in desmids.