New Insights into the Regulation of Stomatal Movements by Red Light, Carbon Dioxide and Circadian Rhythms

Stomata are small adjustable pores formed by pairs of guard cells that enable gas exchange between leaves and the atmosphere, thus directly affecting water loss and CO2 uptake in plants. The current work focuses on the regulation of stomatal movements by red light, carbon dioxide and the circadian system and attempts to uncover molecular mechanisms that control guard cell function. The signaling pathway that underlays stomatal opening in response to red light is yet to be fully elucidated. Here, the HIGH LEAF TEMPERATURE 1 (HT1) protein kinase, known as a negative regulator of high CO2 stomatal closure, is shown to be a key component of stomatal signaling in response to red light (Paper I). It was demonstrated that HT1 is epistatic to the positive regulator of ABAand high CO2induced stomatal closure OPEN STOMATA1 (OST1) protein kinase both in red lightand CO2-induced signal transduction in guard cells (Paper I). A photosynthesis-induced drop in intercellular CO2 as well as processes originating in the photosynthetic electron transport chain (PETC) have been proposed to signal the guard cell response to red light. Investigation of the effect of PETC inhibitors on stomatal conductance in Arabidopsis thaliana ecotypes Col-0 and Ely-1a has suggested the redox state of plastoquinone (PQ) pool to be involved in the regulation of stomatal movements (Paper II). The full mechanisms that link the regulation of stomatal movements to the circadian clock are yet unknown. The blue light receptor, F-box protein and key element of the circadian clock ZEITLUPE (ZTL) was here shown to physically interact with OST1 protein kinase (Paper III). Furthermore, Arabidopsis thaliana mutant plants and Populus transgenic lines that lack the activity of ZTL or OST1 demonstrated similar phenotypes, affected in stomatal movement control (Paper III). The work supports a requirement of both ZTL and OST1 in the regulation of guard cell turgor and suggests a direct link between the circadian clock and OST1 activity.