Stomatal development: new signals and fate determinants

Stomata and pavement cells are produced by a series of asymmetric divisions and progressive fate transitions within a stem cell lineage. In Arabidopsis, this process is regulated so that new lineages can be inserted between previously differentiated cells while maintaining stomatal spacing. The small peptide EPIDERMAL PATTERNING FACTOR 1 may be a positional signal secreted by stomatal precursors to modulate behavior of nearby cells. Signal-receiving cells may use TOO MANY MOUTHS and ERECTA family receptors and a MAPK pathway to regulate initiation of new lineages, promote asymmetric division, and control the plane of spacing divisions. Cell fate transitions are controlled by basic helix–loop–helix transcription factor (bHLH), MYB, and MADS-box transcription factors, and there is evidence of miRNA regulation. These results provide insight into positive and negative influences on stomatal cell transitions and suggest points of potential environmental regulation.