The functional unit for gas exchange between plants and the atmosphere is t
he stomatal complex, an epidermal structure composed of two guard cells, wh
ich delimit a stomatal pore, and their subsidiary cells. In the present wor
k, we define the basic structural unit formed in Arabidopsis thaliana durin
g leaf development, the anisocytic stomatal complex. We perform a cell line
age analysis by transposon excision founding that at least a small percenta
ge of stomatal complexes are unequivocally non-clonal. We also describe the
three-dimensional pattern of stomata in the Arabidopsis leaf. In the epide
rmal plane, subsidiary cells of most stomatal complexes contact the subsidi
ary cells of immediately adjacent complexes. This minimal distance between
stomatal complexes allows each stoma to be circled by a full complement of
subsidiary cells, with which guard cells can exchange water and ions in ord
er to open or to close the pore, In the radial plane, stomata (and their pr
ecursors, the meristemoids) are located at the junctions of several mesophy
ll cells. This meristemoid patterning may be a consequence of signals that
operate along the radial axis of the leaf, which establish meristemoid diff
erentiation precisely at these places. Since stomatal development is basipe
tal, these radially propagated signals may be transmitted in the axial dire
ction, thus guiding stomatal development through the basal end of the leaf.