Transmission electron microscopy was used to study the ultrastructure of epithallial, subepithallial, and initial cells in three species of Lithothamnion: L. glaciale, L. corallioides, and L. sonderi, and four species of Phymatolithon: P. laevigatum, P. purpureum, P. lenormandii, and P. calcareum (Corallinaceae, subfamily Melobesioideae). One of the principal characters used to delimit Lithothamnion is the flared morphology of its epithallial cells. Ultrastructural examination showed that this flared appearance is produced by the trapezoidal outline of the outermost cell wall of the intact epithallial cell, incombination with the presence of the remnant proximal wall of shed epithallial cells. Unlike most coralline species studied ultrastructurally, none of the three species of Lithothamnion had chloroplasts in their epithallial cells. Absence of chloroplasts distinguished epithallial cells from chloroplast-rich initial cells, allowing easy discrimination of these cell types. Extensive wall ingrowths developed on the distal face of initial cells; upon division of an initial cell to produce an epithallial cell and a new initial cell, the ingrowths were inherited by the newly formed epithallial cell. Decalcification of cell walls occurs as part of the process of epithallial cell turnover; subsequent recalcification may then occur. Secondary calcification was present in the wall layer between the remnant proximal wall and the distal epithallial cell surface in L. glaciale and L. corallioides. Lithothamnion corallioides was unique in having an electron-dense hemispherical structure on the distal face of each epithallial cell; this feature may prove to be a useful taxonomic character. In Phymatolithon, the number of epithallial cells per filament varied from none to three. Terminal epithallial cells were domed and much larger than the subterminal epithallial and initial cells. The outer face of the surface epithallial cell was covered by a thick layer of organic material containing the outlines of extracted small crystals of CaCO3, but calcification was very light. Healthy-looking epithallial cells had chloroplasts with thylakoid spacings similar to that of the discoidal chloroplasts of subepithallial cells. Initial cells preparing to divide were surrounded by a thick organic wall. The derivatives of the division were equal in size and until they enlarged, the pair was collectively surrounded by the persistent wall of the old initial cell. The lower cell, the new initial cell, developed distal wall ingrowths, whereas the upper cell, the new epithallial cell, inherited the ingrowths of the parent initial cell. Distal wall ingrowths were still present even in senescent epithallial cells. Epithallial cell morphology has played an important role in the taxonomic separation of Phymatolithon from Lithothamnion. Epithallial fine structure demonstrates additional, systematically useful characters, such as the presence of chloroplasts in the epithallial cells of Phymatolithon and their absence from epithallial cells in Lithothamnion, that are consistent with the recognition of Phymatolilhon. Ultrastructural study also provides new criteria for the proper identification of epithallial and initial cells in Phymatolithon.
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