Expanded roles and divergent regulation of FAMA in Brachypodium and Arabidopsis stomatal development.

Stomata, cellular valves found on the surfaces of aerial plant tissues, present a paradigm for studying cell fate and patterning in plants. A highly conserved core set of related basic helix-loop-helix (bHLH) transcription factors regulates stomatal development across diverse species. We characterized BdFAMA in the temperate grass Brachypodium distachyon and found this late-acting transcription factor was necessary and sufficient for specifying stomatal guard cell fate, and unexpectedly, could also induce the recruitment of subsidiary cells in the absence of its paralogue, BdMUTE. The overlap in function is paralleled by an overlap in expression pattern and by unique regulatory relationships between BdMUTE and BdFAMA. To better appreciate the relationships among the Brachypodium stomatal bHLHs, we used in vivo proteomics in developing leaves and found evidence for multiple shared interaction partners. We reexamined the roles of these genes in Arabidopsis thaliana by testing genetic sufficiency within and across species, and found that while BdFAMA and AtFAMA can rescue stomatal production in Arabidopsis fama and mute mutants, only AtFAMA can specify Brassica-specific myrosin idioblasts. Taken together, our findings refine the current models of stomatal bHLH function and regulatory feedback among paralogues within grasses as well as across the monocot/dicot divide.

Stomatal development in the grasses: lessons from models and crops (and crop models).

When plants emerged from their aquatic origins to colonise land, they needed to avoid desiccation while still enabling gas and water exchange with the environment. The solution was the development of a waxy cuticle interrupted by epidermal pores, known as stomata. Despite the importance of stomata in plant physiology and their contribution to global water and carbon cycles, our knowledge of the genetic basis of stomatal development is limited mostly to the model dicot, Arabidopsis thaliana. This limitation is particularly troublesome when evaluating grasses, whose members represent our most agriculturally significant crops. Grass stomatal development follows a trajectory strikingly different from Arabidopsis and their uniquely shaped four-celled stomatal complexes are especially responsive to environmental inputs. Thus, understanding the development and regulation of these efficient complexes is of particular interest for the purposes of crop engineering. This review focuses on genetic regulation of grass stomatal development and prospects for the future, highlighting discoveries enabled by parallel comparative investigations in cereal crops and related genetic model species such as Brachypodium distachyon.

Grass stomata.

Stomata are adjustable valves through which gas and water exchange occur in plant leaves. Here, McKown and Bergmann highlight the essential function and features of stomata from grasses.