Chloroplast division polarity, a marker of cell division planes during morphogenesis inBangia vermicularis Harvey (Rhodophyceae)

Abstract

Ultrastructural observations of dividing cells inBangia vermicularis revealed a type of chloroplast division (plastokinesis) not previously reported in the red algae. The polarity of this prekaryokinetic process serves as a reliable marker of the plane of cytokinesis, a key factor in establishing thallus morphology. At the onset of division one or more invaginations develop in the envelope of the large, lobed chloroplast and proceed centripetally through the stroma in the plane of the thylakoids, forming narrow cytoplasmic channels (CCs). The thylakoids are realigned somewhat, but are not constricted as the chloroplast is divided into two or more units. The number of resulting chloroplasts and the orientation of the CCs are dependent on cell type. Distinctive cylindrical cells at the base of the filamentous region, immediately distal to the holdfast, are shorter than broad and contain a central nucleus surrounded by a doughnutshaped chloroplast. The cylindrical morphology of the thallus is established early in the first periclinal division as multiple plastokinesis commences, generating several radially-arranged daughter chloroplasts. Cleavage of the original chloroplast is completed during subsequent cell divisions in the initial developmental stage, finally resulting in eight chloroplasts that are distributed to an equal number of wedge-shaped radial cells. Cells distal to the actively dividing basal cells are cuboidal and have a peripheral nucleus. Division of the single chloroplast prior to karyokinesis in these cells results in two or four daughter chloroplasts according to cell type. During or following plastokinesis, multilamellar bodies derived from the CE appear to serve as the source of membranes for the developing septum in the channels. Septa link to proliferations of the plasmalemma in areas of slight cell wall (CW) indentations, and are completed between daughter nuclei after karyokinesis, producing a cleavage channel. Subsequently, primary CW material is deposited between the two septal membranes. The shape and arrangement of daughter cells in each of the developmental stages in the thallus are defined by the planes of cell division. These are indicated by both the orientation of CCs and the polar orientation of nuclear division which is always at right angles to the CC.