Studies on the development of the air pores and air chambers of Marchanda paleacea. II. Ultrastructure of the initial aperture formation with particular reference to cortical microtubule organizing centres

Abstract

The initiation of the intercellular spaces (ISs) of the initial apertures (IAs) follows the formation of surface cavities (SCs). The latter represent slight deepenings in the external periclinal walls of particular superficial thallus cells at the regions where their anticlinal walls meet one another. This event keeps pace with the deposition of wall pads at the wall junctions below the SC. Afterwards, the thickened wall areas are detached and thus the IS is initiated. By an inward development the IS reaches the subprotodermal layer. This is carried out by the coordination of three gradual processes: the inward spreading of the local wall thickening and the following detachment and expansion of the thickened regions. The findings favour the conclusion that the opening of the IS is the outcome of a highly controlled morphogenetic process. The interphase IA cells possess a well-organized cortical microtubule (MT) cytoskeleton, particularly at the area where the IS opens. In these regions, sets of anticlinal and periclinal MTs appear. During the SC stage the anticlinal MTs dominate, while during IS formation, the periclinal ones are most abundant. The above MTs, as well as other ones entering deeper in the cytoplasm, initially converge on the cortical cytoplasm adjacent to the SC and later on the region surrounding the lower part of the growing IS, where vesicles and endoplasmic reticulum are gathered. The observations suggest the continuous function of cortical MT organizing centres in the cytoplasm and (or) the adjoining plasmalemma, initially underneath the SC and later below the lower part of the growing IS, where local wall thickenings are deposited.