Impact of UV radiation on activity of linear furanocoumarins andBacillus thuringiensis var.Kurstaki againstSpodoptera exigua: Implications for tritrophic interactions

Abstract
Acidic fogs with a pH of 2.0 and duration of 2 hr did not reduce the efficacy ofBacillus thuringiensis var.Kurstaki (Berliner). Therefore, the impact of UV radiation was investigated on the interactions between (1) levels of the antibacterial linear furanocoumarins psoralen, bergapten, and xanthotoxin inApium graveolens (L.) occurring following a 2.0 pH acidic fog episode, (2) the noctuidSpodoptera exigua (Hübner), and (3) a sublethal dosage of the microbial pathogenB. thuringiensis var.Kurstaki. Mean time to pupation in the absence of UV radiation (survival was too low to conduct this analysis for insects exposed to UV) was significantly extended by the addition of either psoralens orB. thuringiensis. Larvae developing on diets containingB. thuringiensis plus psoralens required nearly 40% longer to pupate than controls, but their effects were additive as the interaction was not significant. Although the mean times to adult emergence were significantly different, time spent in the pupal stage did not vary significantly between treatments, indicating that increases in larval developmental time were responsible for the observed decrease in developmental rate. Mean time to mortality, a weighted average time of death, was not significantly affected by any of the treatments. In a 2 × 2 × 2 factorial analysis, all main effects (linear furanocoumarins.B. thuringiensis, UV radiation) reduced survival significantly, as did the three-way interaction. Thus, antagonistic interactions with psoralens that would reduce the effectiveness ofB. thuringiensis in the field were not observed. When pairs of main effects were nested within the two levels (presence and absence) of the third factor, several two-way interactions were found. Interestingly, the activity ofB. thuringiensis and the psoralens, individually or in combination, was enhanced by exposure to UV radiation. Implications of this research are discussed for both natural and agricultural ecosystems.