text: There are three major epidermal cell types which all ultimately derive from the outermost (L1) tissue layer of the shoot apical meristem, called protodermal cells: trichomes, pavement cells and guard cells, all of which are arranged in a non-random fashion.
  An asymmetrical cell division occurs in protodermal cells resulting in one large cell that is fated to become a pavement cell and a smaller cell called a meristemoid that will eventually differentiate into the guard cells that surround a stoma. This meristemoid then divides asymmetrically one to three times before differentiating into a guard mother cell. The guard mother cell then makes one symmetrical division, which forms a pair of guard cells. Cell division is inhibited in some cells so there is always at least one cell between stomata.
  Stomatal patterning is controlled by the interaction of many signal transduction components such as EPF (Epidermal Patterning Factor), ERL (ERecta Like) and YODA (a putative MAP kinase kinase kinase). Mutations in any one of the genes which encode these factors may alter the development of stomata in the epidermis. For example, a mutation in one gene causes more stomata that are clustered together, hence is called Too Many Mouths (TMM). Whereas, disruption of the SPCH (SPeecCHless) gene prevents stomatal development all together.  Inhibition of stomatal production can occur by the activation of EPF1, which activates TMM/ERL, which together activate YODA. YODA inhibits SPCH, causing SPCH activity to decrease, preventing asymmetrical cell division that initiates stomata formation. Stomatal development is also coordinated by the cellular peptide signal called stomagen, which signals the activation of the SPCH, resulting in increased number of stomata.
  Environmental and hormonal factors can affect stomatal development. Light increases stomatal development in plants; while, plants grown in the dark have a lower amount of stomata. Auxin represses stomatal development by affecting their development at the receptor level like the ERL and TMM receptors. However, a low concentration of auxin allows for equal division of a guard mother cell and increases the chance of producing guard cells.
  Most angiosperm trees have stomata only on their lower leaf surface. Poplars and willows have them on both surfaces. When leaves develop stomata on both leaf surfaces, the stomata on the lower surface tend to be larger and more numerous, but there can be a great degree of variation in size and frequency about species and genotypes. White ash and white birch leaves had fewer stomata but larger in size. On the other hand sugar maple and silver maple had small stomata that were more numerous.