This light- and electron-microscope study of four species of Sphagnum
reveals that stem elongation involves
meristematic activities unique to the group and hitherto unrecognized.
The internal tissue of the mature stem
arises by the concerted activity of an apical (primary) and a subapical
(secondary) meristem. The primary
meristem comprises the immediate derivatives of the single apical cell.
Following a small number of divisions, the
primary derivatives differentiate into highly vacuolate parenchymatous
cells with a storied arrangement.
Subsequently, the large vacuoles are replaced by numerous small vacuoles
and the cells then divide repeatedly, by
transverse septa, producing files of about nine short cells. Finally, ninefold
elongation of these secondary cells is
responsible for extension growth of the main stem below the mature capitulum.
An early step in primary
differentiation is the confinement of pre-existing plasmodesmata to distinct
pitted areas. Further enlargement of
the cells during primary and secondary differentiation involves the thickening
of non-pitted wall areas, followed
by expansion and thinning out, while the pitted areas remain virtually
unchanged. A cortical array of microtubules
is regularly found in association with non-pitted wall areas, while the
unexpanded pitted areas are associated with
smooth endoplasmic reticulum showing continuity with desmotubules. Though
sharing much the same cytology
as the conducting cells in bryoid mosses, in terms of their development
the central stem cells in Sphagnum are not
homologous with those of other mosses. The unique mode of stem development
may be an important factor in the
ecological success of Sphagnum.