Influence of environmental factors on stomatal development.

Stomata play a pivotal role in the regulation of gas exchange in flowering plants and are distributed throughout the aerial epidermis. In leaves, the pattern of stomatal distribution is highly variable between species but is regulated by a mechanism that maintains a minimum of one cell spacing between stomata. In Arabidopsis, a number of the genetic components of this mechanism have been identified and include, SDD1, EPF1 and the putative receptors TMM and the ERECTA-gene family. A mitogen-activated protein (MAP) kinase signalling cascade is believed to act downstream of these putative receptors while a number of transcription factors including SPCH, MUTE and FAMA have been identified that control consecutive steps of stomatal development. The environment also has significant effects on stomatal development. In a number of species both light intensity and CO(2) concentrations have been shown to influence the frequency at which stomata develop on leaves. Long-distance signalling mechanisms have been implicated in these environmental responses with the conditions sensed by mature leaves determining the stomatal frequency in developing leaves. Thus, changes in the environment appear to act by modulating the developmental and patterning pathways to determine stomatal frequency.