Studies on the formation of 'floating' guard cell mother cells in Anemia.

The protodermal cells producing the 'floating' guard cell mother cells (GMCs) in three Anemia species undergo an extraordinary polarization and an unexpected shaping. During interphase an intercellular space is initiated at the internal proximal end of the cell, while the polar region bulges outwards. At this stage a microtubule girdle traverses the cortical cytoplasm underneath the rims of the external periclinal wall curvature. In addition, another system of microtubules converges on a cortical site adjoining the wall delimiting the intercellular space and, or, the neighbouring region of the internal periclinal wall (internal polar cortical site, IPCS). Vacuoles are found in all regions of the cell except for that between the centrally located nucleus and the intercellular space. As the cell approaches mitosis, the growing vacuolar system retreats from the cytoplasmic region below the external periclinal wall curvature. In most cells the polarized cytoplasm forms an inclined truncated cone, the bases of which abut on the external periclinal wall curvature and the wall lining the IPCS. The organization of the cortical microtubule cytoskeleton does not change significantly during preprophase-prophase. A preprophase microtubule band (PMB) is localized in the cortex lining the rims of the external periclinal wall curvature, while some microtubules traverse the IPCS and the cytoplasm adjacent to the neighbouring wall regions. The mitotic spindle axis is diagonal, while the cell plate separating the GMCs exhibits an unusual mode of growth. It gradually encircles the proximal daughter nucleus, becoming funnel-shaped. One of its periclinal edges fuses with the external periclinal wall area lined by the PMB cortical zone and the other with the internal periclinal wall area adjoining the IPCS. The latter region seems to behave like the PMB cortical zone. The results show that the morphogenetic mechanism underlying the formation of the conical GMCs includes a series of highly integrated processes, initiated or carried out during cell polarization.