Flooding and Herbivory Interact to Alter Volatile Organic Compound Emissions in Two Maize Hybrids

Flooding is a major plant abiotic stress factor that is frequently experienced by plants simultaneously with other biotic stresses, including herbivory. How plant volatile emissions, which mediate interactions with a wide range of organisms, are influenced by flooding and by multiple co-occurring stress factors remains largely unexplored. Using Spodoptera frugiperda (Lepidoptera: Noctuidae) (fall armyworm) as the insect pest and two maize (Zea mays, L. Poaceae) hybrids differentially marketed for conventional and organic production, we assessed the effects of flooding, herbivory, and both stress factors on the composition of blends of emitted volatiles. Headspace volatiles were collected from all treatment combinations seven days after flooding. We documented metrics indicative of biomass allocation to determine the effects of individual and combined stressors on plant growth. We also evaluated relationships between volatile emissions and indicators of soil chemical characteristics as influenced by treatment factors. Flooding and herbivory induced the emission of volatile organic compounds (VOCs) in similar ways on both maize hybrids, but the interaction of both stress factors produced significantly larger quantities of emitted volatiles. Thirty-eight volatile compounds were identified, including green leaf volatiles, monoterpenes, an aldehyde, a benzoate ester, sesquiterpenes, a diterpene alcohol, and alkane hydrocarbons. The hybrid marketed for organic production was a stronger VOC emitter. As expected, plant biomass was detrimentally affected by flooding. Soil chemical properties were less responsive to the treatment factors. Taken together, the results suggest that flooding stress and the interactions of flooding and insect attack can shape the emission of plant volatiles and further influence insect-plant interactions.