Richard A. Werner

Effect of nitrogen fertilization upon the secondary chemistry and nutritional value of quaking aspen (Populus tremuloides Michx.) leaves for the large aspen tortrix (Choristoneura conflictana (Walker))

SummaryWe investigated the effects of nitrogen fertilization upon the concentrations of nitrogen, condensed tannin and phenolic glycosides of young quaking aspen (Populus tremuloides) leaves and the quality of these leaves as food for larvae of the large aspen tortrix (Choristoneura conflictana), a Lepidopteran that periodically defoliates quaking aspen growing in North America. Nitrogen fertilization resulted in decreased concentrations of condensed tannin and phenolic glycosides in aspen leaves and an increase in their nitrogen concentration and value as food for the large aspen tortrix. These results indicate that plant carbon/nutrient balance influences the quality of aspen leaves as food for the large aspen tortrix in two ways, by increasing the concentrations of positive factors (e.g. nitrogen) and decreasing the concentrations of negative factors (eg. carbon-based secondary metabolites) in leaves. Addition of purified aspen leaf condensed tannin and a methanol extract of young aspen leaves that contained condensed tannin and phenolic glycosides to artificial diets at high and low levels of dietary nitrogen supported this hypothesis. Increasing dietary nitrogen increased larval growth whereas increasing the concentrations of condensed tannin and phenolic glycosides decreased growth. Additionally, the methanol extract prevented pupation. These results indicate that future studies of woody plant/insect defoliator interactions must consider plant carbon/nutrient balance as a potentially important control over the nutritional value of foliage for insect herbivores.

Chemical model for short-term induction in quaking aspen (Populus tremuloides) foliage against herbivores

Simulated large aspen tortrix (Choristoneura conflictana) herbivory of quaking aspen (Populus tremuloides) induces significant increases in concentrations of two phenol glycosides, salicortin and tremulacin, in leaves within 24 hr. Crushing of leaf tissue, as must occur when aspen leaves are eaten by chewing insects such as the large aspen tortrix, results in conversion of salicortin and tremulacin to 6-hydroxy-2-cyclohexenone (6-HCH). Salicortin, tremulacin, 6-HCH, and its degradation product, catechol, are all toxic to the large aspen tortrix when fed on an artificial diet. These damage-induced chemical changes provide a plausible mechanism for short-term resistance induced in aspen leaves by insect herbivory.

Effects of Mineral Nutrition on Delayed Inducible Resistance in Alaska Paper Birch

In subarctic forests, birch (Betula) trees respond to severe (50-100%) manual defoliation by delayed inducible resistance (DIR). This plant response to defoliation is characterized by a decline in the nutritional quality of leaves for immature insects for several years after defoliation events, and concomitant changes in leaf chemistry that may be detrimental to insect nutrition, that is, a decline in leaf nitrogen and an increase in leaf phenols. Two explanations of delayed inducible resistance have been proposed. (1) The active defense response hypothesis claims that delayed inducible resistance is an active response to defoliation per se rather than merely a passive consequence of recovery from the stress of severe defoliation. (2) In contrast, the carbon-nutrient balance (CNB) hy- pothesis claims that delayed inducible resistance is caused by nutritional stress resulting from severe defoliation. We used two experiments to test these hypotheses. (1) In a three- way factorial field experiment we reared spear-marked black moth (Rheumaptera hastata) larvae on Alaska paper birch (B. resinifera) saplings that had experienced combinations of 100% manual defoliation and fertilization with N and P in previous years, and measured larval survival and pupal mass. In association with these measurements of larval perfor- mance, we assayed leaf condensed tannin levels and the concentrations of N and P in leaves, and correlated the results of these assays with larval performance. (2) In a laboratory experiment we tested the biological activity of condensed tannin and linalool, the major secondary metabolites of Alaska paper birch leaves, by treating leaves collected from previously undefoliated Alaska paper birch saplings with combinations of condensed tannin and linalool, and measuring the performance of spear-marked black moth larvae reared on these leaves. Our results supported predictions of the carbon-nutrient balance hypoth- esis. We found that fertilization with nitrogen, the nutrient limiting growth of Alaska paper birch in our study site, mitigated delayed inducible resistance, and that condensed tannin is likely to be the major chemical cause of delayed inducible resistance in Alaska paper birch. In our field experiment we also found that fertilization of Alaska paper birch with phosphorus, a nutrient that does not limit the growth of Alaska paper birch in our study site, affected levels of condensed tannin in leaves and the saplings defoliation history influenced this effect. Thus, future studies of the effects of mineral nutrition on secondary metabolite production by woody plants and their responses to herbivory should consider

Toxicity of Carbaryl toward the Southern Pine Beetle in Filter Paper, Bark and Cut Bolt Bioassays

Loblolly pine bolts sprayed with 2% carbaryl (Sevimol® 40% A. I. Flowable) and 1% chlorpyrifos were exposed to southern pine beetle, Dendroctonus frontalis Zimmermann. Chlorpyrifos treatment effectively prevented attack by southern pine beetle; however, the bolts were not protected by 2% carbaryl. For the carbaryl-treated bolts, the number of egg niches and larval mines were significantly reduced. However, the number of adult beetles entering the host, the length of the parent galleries, the number of pupal chambers, brood adults, and emergence holes were not significantly (P = 0.05) different from untreated control bolts. A bark surface assay indicated that 2% carbaryl killed < 50% of the beetles at 24 h. Filter paper assay showed that the LC50 value was 0.07% and 0.01% at 24 and 48 h, respectively.