Stomatal Patterning

Stomatal development is a model for understanding the integration of multiple inputs into a simple cell fate choice. A small but growing network of genes has been identified that are responsible for stomatal development in the model plant Arabidopsis: a set of transcription factors control the final stages of guard cells production and proteins suspected to play a role in cell–cell communication affect how many stomata are formed and their distinct distributions in various organs.

Keywords: stomata; guard cells; receptor-like kinase; cell signalling; development

Figure 1. Stomatal structure and development. (a) Photomicrograph of typical stoma. Black arrow points to paired guard cells and black arrowhead indicates subsidiary cells (b) Source of environmental inputs into stomatal development (c) Diagram of stages in stomatal development. Cells undergo entry divisions to create meristemoids (M, light blue). Meristemoids either differentiate into guard mother cells (GMC, dark blue) or continue amplifying divisions. GMCs differentiate into guard cells (GC, violet). Spacing divisions take place in cells adjacent to stomatal lineage cells.
Figure 2. Diversity of stomatal pattern. (a) Scanning electron micrograph (SEM) image of pine needle, white arrow points to one of many stomata arranged in long files. (b–c) Resin impression of a leaf of a monocot, the ‘spider plant’ (b) bottom of leaf, white arrow points to one stoma within a highly regular array of stomata positioned between the leaf veins (c) top of the same leaf lacking any stomata. (d–e) SEM of the top (d) and bottom (e) of a California poppy (dicot) leaf, white arrows point to stomata present on both surfaces (f) impression of a developing Arabidopsis leaf epidermis. Arrows in inset (at higher magnification) show stomata obeying the one-cell spacing rule.
Figure 3. Model of potential molecular signalling pathway for Arabidopsis stomatal development. Cells in the stomatal lineage produce a mobile signal by the cleavage activity of SDD1. The peptide signal is perceived at the membrane of the responding cell by the leucine-rich repeat containing protein, TMM and the ERL family protein kinases. The exact nature of the interaction between TMM and the ERL family is complex and has not been fully elucidated. Each of the four proteins could potentially dimerize with each of the others, and it is not known whether ligand binding induces dimerization, or whether dimers are required to perceive ligand. Upon perception by the receptors, the signal is then transduced via an unknown mechanism to a MAP kinase cascade headed by YODA. The cascade ultimately phosphorylates unknown downstream targets and results in a repression of the stomatal fate.
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 References
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 Further Reading
    Bergmann DC (2004) Integrating signals in stomatal development. Current Opinion in Plant Biology 7: 26–32.
    Ingram GC (2005) Plant development: spacing out stomatal pores. Current Biology 15: R663–R665.
    Larkin JC, Brown ML and Schiefelbein J (2003) How do cells know what they want to be when they grow up? Lessons from epidermal patterning in Arabidopsis. Annual Review of Plant Biology 54: 403–430.
    Leonhardt N, Kwak JM, Robert N et al. (2004) Microarray expression analyses of Arabidopsis guard cells and isolation of a recessive abscisic acid hypersensitive protein phosphatase 2C mutant. Plant Cell 16: 596–615.
    Masle J, Gilmore SR and Farquhar GD (2005) The ERECTA gene regulates plant transpiration efficiency in Arabidopsis. Nature 436: 866–870.
    book Nadeau JA and Sack FD (2002) "Stomatal development in Arabidopsis". In: Somerville C and Meyerowitz E (eds) The Arabidopsis Book, pp. 1–28. Rockville, MD: American Society of Plant Biologists
    Yang M and Sack FD (1995) The too many mouths and four lips mutations affect stomatal production in Arabidopsis. Plant Cell 7: 2227–2239.
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MacAlister, Cora A, and Bergmann, Dominique C(Jan 2007) Stomatal Patterning. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0020125]