SPEECHLESS and MUTE Mediate Feedback Regulation of Signal Transduction during Stomatal Development

Abstract

Stomatal density, spacing, and patterning greatly influence the efficiency of gas exchange, photosynthesis, and water economy. They are regulated by a complex of extracellular and intracellular factors through the signaling pathways. After binding the extracellular epidermal patterning factor 1 (EPF1) and 2 (EPF2) as ligands, the receptor-ligand complexes activate by phosphorylation through the MAP-kinase cascades, regulating basic helix-loop-helix (bHLH) transcription factors SPEECHLESS (SPCH), MUTE, and FAMA. In this review, we summarize the molecular mechanisms and signal transduction pathways running within the transition of the protodermal cell into a pair of guard cells with a space (aperture) between them, called a stoma, comprising asymmetric and symmetric cell divisions and draw several functional models. The feedback mechanisms involving the bHLH factors SPCH and MUTE are not fully recognized yet. We show the feedback mechanisms driven by SPCH and MUTE in the regulation of EPF2 and the ERECTA family. Intersections of the molecular mechanisms for fate determination of stomatal lineage cells with the role of core cell cycle-related genes and stabilization of SPCH and MUTE are also reported.

LEA13 and LEA30 Are Involved in Tolerance to Water Stress and Stomata Density in Arabidopsis thaliana

“…SPCH regulates the meristemoid mother cell (MMC) transition and asymmetric cell divisions. MUTE regulates meristemoid transition in the guard mother cell (GMC), while FAMA regulates asymmetric divisions that give rise to guard cells (GC) [ 6 ]. Additionally, stomatal density and pattern are determined by signals involving the EPIDERMAL PATTERNING FACTORS (EPFs) family, plant hormones, and environmental clues.…”