An Abscisic Acid-Independent Oxylipin Pathway Controls Stomatal Closure and Immune Defense in Arabidopsis
Stomata are microscopic pores that are present in the epidermis of the aerial parts of higher plants, such as the leaves. These pores, which are flanked by a pair of cells called guard cells, regulate transpiration and the exchange of gas between leaves and the atmosphere. It is well documented that the phytohormone abscisic acid (ABA) is a key regulator that controls the osmotic pressure in guard cells, allowing pore size to be adjusted in response to environmental conditions. Recently, stomata have also been shown to play an important role in the innate immune response. Indeed, upon contact with microbes, plants actively close stomata to prevent the entry of microbes and the consequent colonization of host tissue. This response is known as the stomatal defense response. However, the molecular mechanisms that regulate this defense response are not well understood. Using a variety of approaches, we show in this study that LOX1, a gene that encodes lipoxygenase (LOX) in guard cells, plays a major role in stomatal defense in the model plant Arabidopsis thaliana. Mutations in LOX1 impair stomatal closure and make plants more susceptible to the bacterium Pseudomonas syringae pv. tomato. We also show that several LOX-derived metabolites, the oxylipins, are potent inducers of stomatal closure. Finally, we provide evidence to show that ABA plays only a minor role in stomatal defense response, specifically by modulating this response.