1 Host plant terpenes can influence attraction of conifer bark beetles to their aggregation pheromones: both synergistic and inhibitory compounds have been reported. However, we know little about how varying concentrations of individual monoterpenes affect responses. 2 We tested a gradient of ratios of a-pinene, the predominant monoterpene in host pines in the Great Lakes region of North America, to Ips pini's pheromone, racemic ipsdienol plus lanierone. 3 Ips pini demonstrated a parabolic response, in which low concentrations of a-pinene had no effect on attraction to its pheromone, intermediate concentrations were synergistic and high concentrations were inhibitory. These results suggest optimal release rates for population monitoring and suppression programmes. 4 Inhibition of bark beetle attraction to pheromones may be an important component of conifer defences. At terpene to pheromone ratios emulating emissions from trees actively responding to a first attack, arrival of flying beetles was low. This may constitute an additional defensive role of terpenes, which are also toxic to bark beetles at high concentrations. 5 Reduced attraction to a low ratio of a-pinene to pheromone, as occurs when colonization densities become high and the tree's resin is largely depleted, might reflect a mechanism for preventing excessive crowding. 6 Thanasimus dubius, the predominant predator of I. pini, was also attracted to ipsdienol plus lanierone, but its response differed from that of its prey. Attraction increased across all concentrations of a-pinene. This indicates that separate lures are needed to sample both predators and bark beetles effectively. It also provides an opportunity for maximizing pest removal while reducing adverse effects on beneficial species. This disparity further illustrates the complexity confronting natural enemies that track chemical signals to locate herbivores.

Salivary glucose oxidase (GOX) has been reported in a few insect species where it plays a role in protection against infectious disease. Our recent research has focused on the role of this salivary enzyme in the noctuid Helicoverpa zea, where it functions as an effector to suppress the induced defenses of the host plant Nicotiana tabacum. In this study, we examined the labial gland GOX activities in 23 families of Lepidoptera (85 species) and two families of plant-feeding Hymenoptera (three species). We analyzed the relationship between host breadth and GOX activities, and we found a significant relationship, where highly polyphagous species were more likely to possess relatively high levels of GOX compared to species with more limited host range. We also examined the effect of diet on GOX activity and found that the host plant had a significant effect on enzyme activity. The significance of these findings is discussed in relation to caterpillar host breadth.

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Chemical communication by many insect species involves complex signals of both insect and plant origin. Much attention has been focused on the behavioral activities of these components but less on their sources of variation, despite implications for evolutionary theory and pest management. We studied variation in chemical signaling at host, tree-within-host, and beetle-on-tree scales using tunneling male pine engravers [Ips pini (Say)] on jack, Pinus banksiana Lamb, red, P. resinosa Aiton, and white, P. strobus L. pines. Pine engravers are distributed transcontinentally, and stereoisomeric ratios of their principal pheromone component ipsdienol varies regionally. Linear mixed-effects models were used to examine variation in monoterpene and pheromone volatile profiles, determined by gas chromatography. Phloem from white pine had the greatest concentration of monoterpenes, although insects tunneling in white pine produced the smallest ratios of monoterpenes to pheromones (1:2) in their volatile plumes relative to jack and red pine (1:1). Beetle-to-beetle variation in plume composition was approximately 2-9 times greater than the inter-tree variation within a tree species. The stereoisomeric ratio of ipsdienol was highly consistent within the pheromone component of the plume. The little variation present existed almost entirely at the level of the insects. Within the pheromone component of the plume in a given host species, there was up to 13 times more beetle-to-beetle than tree-to-tree variation. This magnitude was almost double the magnitudes of the ratios among components within the entire plumes. Implications to the behavioral ecology of bark beetle communication, such as potential strategies of cheating and predator avoidance, are discussed.

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Conifers comprise a substantial portion of the host range of the gypsy moth, Lymantria dispar L. In this study, the effects and fates of the predominant conifer phytochemicals, monoterpenes and diterpene acids, were determined. Gypsy moth larvae were fed physiological levels of three monoterpenes (bornyl acetate, limonene, and myrcene) and two diterpene acids (isopimaric acid and neoabietic acid) for the duration of their second, third, and fourth stadia. Frass was collected daily, and larvae and exuviae were collected after the completion of the stadium. These samples were chemically analyzed for parent terpenes or their metabolites. Overall, gypsy moth larvae were highly tolerant of all terpenes administered, despite some treatment effects. Limonene, myrcene, isopimaric acid, and neoabietic acid were not recovered suggesting that, once ingested, these terpenes were converted into other products. Bornyl acetate was metabolized in part to borneol and also was recovered in small quantities from frass. Its metabolite borneol was mostly excreted, with a small amount remaining in body tissues. Additional experiments were performed, determining that monoterpenes were effectively ingested, and that monoterpenes were not lost substantially through volatilization from diet or larvae. There was no evidence that terpene metabolism was due to P450 enzymes or esterases, although these results are inconclusive. Gut bacteria do not appear to be primarily responsible for terpene metabolism.

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