The role of strigolactones in regulation of stomatal conductance and plant-pathogen interactions inArabidopsis thaliana

AbstractStrigolactones are a group of phytohormones that control shoot branching inArabidopsis thaliana. However, in recent years they have been shown to affect many other plant processes. We previously showed that the strigolactone perception mutantmore axillary branches 2 (max2)has increased susceptibility to plant pathogenic bacteria as a result of more open stomata as well as alterations in hormonal signalling. Here we show that both, strigolactone biosynthesis- (max3andmax4), and perception mutants (max2anddwarf14) are significantly more sensitive toPseudomonas syringaeDC3000. Moreover, in response toP. syringaeinfection, high levels of SA accumulated inmax2and this mutant was ozone sensitive. To search for the mechanisms that could explain pathogen- and ozone sensitivity we performed gene expression analysis and several different assays that explore the function of guard cells and regulation of guard cell signalling.Treatments with GR24 (a strigolactone analogue) resulted in very modest changes in defence-related gene expression. In contrast, guard cell function was clearly impaired inmax2and depending on the assay used, also inmax3, max4andd14mutants. Moreover, stomatal responses to stimuli that cause stomatal closure in wild-type plants (darkness, high CO2and ABA) were analysed in the strigolactone mutants. In darkness both strigolactone biosynthesis and perception mutants showed reduced stomatal closure, whereas the response to high CO2was impaired only inmax2andd14. The response to ABA was not impaired in any of the mutants. To position the role of MAX2 in the guard cell signalling network,max2was crossed with mutants defective in ABA biosynthesis (aba2), in guard cell ABA signalling (ost1) and a scaffold protein required for proper ion channel activity (ghr1). The stomatal conductance of double mutants was consistently higher than the corresponding single mutants, suggesting that MAX2 acts in a signalling pathway that functions in parallel to the well characterized guard cell ABA signalling pathway. We propose that the impaired defence responses ofmax2is related to more open stomata that allows increased entry of bacteria or air pollutants like ozone. Furthermore, as MAX2 appears to act in a specific branch of guard cell signalling (related to CO2signalling), this protein could be one of the elusive components that allow guard cells to distinguish between different environmental conditions.