STOMATAL ONTOGENY/ DEVELOPMENT

Stomata begin to develop in a leaf shortly before the main period of meristematic
activity in the epidermis is completed and continue to arise through a considerable
part of the later expansion of the leaf by cell enlargement. Stomatal development
begins with an asymmetric, or unequal, anticlinal division of a protodermal cell. This
division results in two cells, one that is usually larger and resembles the other
protodermal cells and a second that is usually markedly smaller and contains
densely staining cytoplasm and a large nucleus. The smaller of these two cells is
called the stomatal meristemoid. In some species the sister cell of the
meristemoid may divide asymmetrically again and give rise to another meristemoid.
Depending on the species, the meristemoid may function directly as the guard
mother cell (guard-cell mother cell, stoma mother cell) or give rise to the guard
mother cell after further divisions. The formation of the stomatal complex requires
migration of the nucleus to specific sites in the parent cells before cell division and
precise placement of the division planes. An equal division of the guard mother cell
gives rise to the two guard cells, which through differential wall deposition and
expansion acquire their characteristic shape. The middle lamella at the site of the
future pore swells, and the connection between the cells is weakened there. The
cells then separate at the site and thus the stomatal opening is formed. The exact
cause(s) of separation of the ventral walls at the pore site has not been identified,
but three possibilities have been considered: enzymic hydrolysis of the middle
lamella, tension brought about by increase in guard cell turgor, and formation of the
cuticle, which eventually lines the newly formed pores (Sack, 1987).
As the development proceeds, subsidiary cells or neighboring cells may arise from
the same meristemoid as the stoma or from cells that are not directly related
ontogenetically with the guard mother cell. On this basis three major categories of
stomatal ontogeny have been recognized (Pant, 1965)

If cells of the stoma and subsidiary cells are derived from the same mother
cell, is referred to as mesogenous development. It is characteristic of
many gymnosperms, in which this pattern of development is also termed
syndetocheilic.

If the guard cells and the subsidiary cells are derived from different
protodermal initials, development is referred to as perigenous. This pattern
is common in many dicotyledons. In gymnosperms this pattern of
development is also called haplocheilic.

In some plants in which one of two subsidiary cells has the same origin as
the guard cells, development is termed mesoperigenous.