Take a deep breath: peptide signalling in stomatal patterning and differentiation.

Stomata are pores in the leaf surface that open and close to regulate gas exchange and minimize water loss. In Arabidopsis, a pair of guard cells surrounds each stoma and they are derived from precursors distributed in an organized pattern on the epidermis. Stomatal differentiation follows a well-defined developmental programme, regulated by stomatal lineage-specific basic helix-loop-helix transcription factors, and stomata are consistently separated by at least one epidermal cell (referred to as the 'one-cell-spacing rule') to allow for proper opening and closure of the stomatal aperture. Peptide signalling is involved in regulating stomatal differentiation and in enforcing the one-cell-spacing rule. The cysteine-rich peptides EPIDERMAL PATTERNING FACTOR 1 (EPF1) and EPF2 negatively regulate stomatal differentiation in cells adjacent to stomatal precursors, while STOMAGEN/EPFL9 is expressed in the mesophyll of developing leaves and positively regulates stomatal development. These peptides work co-ordinately with the ERECTA family of leucine-rich repeat (LRR) receptor-like kinases and the LRR receptor-like protein TOO MANY MOUTHS. Recently, specific ligand-receptor pairs were identified that function at two different stages of stomatal development to restrict entry into the stomatal lineage, and later to orient precursor division away from existing stomata. These studies have provided the groundwork to begin to understand the molecular mechanisms involved in cell-cell communication during stomatal development.