Foliage constituents of douglas fir (Pseudotsuga menziesii (Mirb.) Franco (Pinaceae)): Their seasonal variation and potential role in douglas fir resistance and silviculture management.

Seasonal changes in the production of primary nutrients (soluble carbohydrates) and secondary metabolites (terpenes, monomeric phenolics, and tannins) in the current needle tissue of Douglas fir were investigated. All four classes of compounds showed significant seasonal changes in concentration during foliage development. Most terpenes increased significantly in concentration from June 11 to August 3, and then showed declining concentrations to September 20. The most dramatic and significant seasonal increases occurred inα-pinene, camphene, and bornyl acetate concentrations. The monomeric phenolics chlorogenic acid, taxifolin glucoside, quercetin galactoside, and those unknown phenolics showed an overall trend of declining in concentration from June 11 through September 20. However, considerable variation between sampling dates in the concentration of these phenolics was noted. Tannin concentration decreased significantly from June 11 to July 9, and then increased in concentration to the September 20 sampling date. Fructose, galactose, glucose, and sucrose tended to decrease from June 11 to September 20. However, significant variation between sampling dates was evident in these compounds as well. Galactose was the major compound in the soluble carbohydrate fraction, amounting to almost 80% of the total concentration throughout the growing season. These data suggest that if phenolics and tannins function as defenses, they would only affect second- and possibly third-instar budworm larvae during the time that these instars mine the buds. Camphene,α-pinene, and bornyl acetate increased in concentration throughout the growing season and may be effective deterrents to the budworm. Both bornyl acetate and camphene have been shown in field and laboratory studies to increase larval mortality and adversely affect budworm larval growth. Carbohydrates generally act as nutrients that enhance herbivore growth. However, in a previous study, galactose was found to cause reduced budworm larval growth and increased larval mortality.