CELL DIVISION IN BULBOCHAETE. II. HAIR CELL FORMATION1

Most vegetative cells of Bulbochaete, and all those of Oedogonium, possess an apical, circular discontinuity in the structure of their secondary wall. Rupture of the wall at this precise site permits expansion of the ring during cell division and release of the zoospore following zoosporogenesis. Certain cells of Bulbochaete (always the apical daughter cell of a division pair) lack this type of discontinuity. Instead, the apical wall is thinned out on one side, so that the cell bulges asymmetrically. In the middle of the bulge is a wall discontinuity which extends only part way around the cell. The wall will rupture here, too, for zoospore release, but if a cell having such a wall, divides, it invariably does so asymmetrically, with one pole of the spindle located in the bulge. Cytokinesis then cuts off a small, colorless daughter cell. The wall ruptures at the discontinuity, and this daughter cell emerges through the slit and differentiates into a hair. The creation of hairs in such cells commences with the deposition of a pad of primary wall lining the bulge. Golgi bodies are involved in its secretion, but not in that of a secondary wall layer which forms next in the premitotic cell and covers the primary wall. The cell becomes polarized; the nucleus migrates toward this region as the chloroplast moves aside. After the asymmetric mitosis, a curved phycoplast cuts off the hair cell nucleus and prevents the chloroplast from moving back into the future hair, whose cytoplasm soon loses much of its affinity for heavy metal stains. Following rupture of the parental wall, the phycoplast moves some distance past the limits of the newly deposited secondary wall layer and then forms a cross wall under the hair. The secondary wall of the hair is not continuous with the secondary wall structure of the parental cell; the circular discontinuity that arises around the base of the bulging parental wall is then perpetuated and accentuated as the hair's secondary wall thickens. This wall weakening becomes the dislocation that will predetermine the site of the ring and consequently the direction of cell expansion in the next normal division of the cell subtending the hair. Abnormal ring formation and the creation of terminal twin hairs have also been examined. The lip of the growing hair contains a characteristic organization of membranes and other components which may be related to the organization of the hair's numerous longitudinally oriented microtubules. These results are discussed in terms of the morphology of the wall in the Oedogionales generally. The creation of the special wall morphology that leads to hair cell formation is considered to be ontogenetically related to a similar wall morphology that is involved in formation of the fertilization pore of the oogonium.