Stomata: gatekeepers of uptake and defense signaling by green leaf volatiles in maize

Plants adapt to balance growth-defense tradeoffs in response to both biotic and abiotic stresses. Green leaf volatiles (GLVs) are released after biotic and abiotic stresses and function as damage-associated signals in plants. Although, GLVs enter plants primarily through stomata, the role of stomatal regulation on the kinetics of GLV uptake remains largely unknown. Here, we illustrate the effect of stomatal closure on the timing and magnitude of GLV uptake. We closed stomata by either exposing maize (Zea mays) plants to darkness or applying abscisic acid, a phytohormone that closes the stomata in light. Then, we exposed maize seedlings to (Z)-3-hexen-1-ol and compared its dynamic uptake under different stomatal conditions. Additionally, we used (E)-3-hexen-1-ol, an isomer of (Z)-3-hexen-1-ol not made by maize, to exclude the role of internal GLVs in our assays. We demonstrate that closed stomata effectively prevent GLV entry into exposed plants, even at high concentrations. Furthermore, our findings indicate that reduced GLV uptake impairs GLV-driven induction of biosynthesis of sesquiterpenes, a group of GLV-inducible secondary metabolites, with or without herbivory. These results elucidate how stomata regulate the perception of GLV signals, thereby dramatically changing the plant responses to herbivory, particularly under water stress or dark conditions.