The initiation of stomata, microscopic valves in the epidermis of higher plants that control of gas exchange, requires a co-ordinated sequence of asymmetric and symmetric divisions, which is under tight environmental and developmental control. Arabidopsis leaves grown under elevated photosynthetic photon flux density have a higher density of stomata. STOMAGEN encodes an epidermal patterning factor produced in the mesophyll, and our observations indicated that elevated photosynthetic irradiation stimulates STOMAGEN expression. Our analysis of gain and loss of function of STOMAGEN further detailed its function as a positive regulator of stomatal formation on both sides of the leaf, not only in terms of stomatal density across the leaf surface but also in terms of their stomatal index. STOMAGEN function was rate limiting for the light response of the stomatal lineage in the adaxial epidermis. Mutants in pathways that regulate stomatal spacing in the epidermis and have elevated stomatal density, such as stomatal density and distribution (sdd1) and too many mouth alleles, displayed elevated STOMAGEN expression, suggesting that STOMAGEN is either under the direct control of these pathways or is indirectly affected by stomatal patterning, suggestive of a feedback mechanism. These observations support a model in which changes in levels of light irradiation are perceived in the mesophyll and control the production of stomata in the epidermis by mesophyll-produced STOMAGEN, and whereby, conversely, stomatal patterning, either directly or indirectly, influences STOMAGEN levels.

Cardiff School of Biosciences The Sir Martin Evans Building Museum Avenue Cardiff CF10 3AX UK

Department of Experimental Plant Biology Faculty of Science University of South Bohemia Branisovska 1760 370 05 Ceske Budejovice Czech Republic

Institute of Experimental Botany The Czech Academy of Sciences Rozvojová 263 165 02 Prague 6 Czech Republic

Institute of Plant Molecular Biology The Biology Centre of the Czech Academy of Sciences Branisovska 31 370 05 Ceske Budejovice Czech Republic

Institute of Plant Molecular Biology The Biology Centre of the Czech Academy of Sciences Branisovska 31 370 05 Ceske Budejovice Czech Republic Department of Experimental Plant Biology Faculty of Science University of South Bohemia Branisovska 1760 370 05 Ceske Budejovice Czech Republic

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