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Monoterpenes, sesquiterpenes, and phenolic and flavonoid glycosides typical of the current year's foliage of Douglas fir (Pseudotsuga menziesii) were bioassayed using agar diets to determine the effect of these compounds on natural and colony populations of western spruce budworm (Choristoneura occidentalis). Several terpenes adversely affected budworm larval growth, supporting previous field and laboratory studies. However, larvae collected from a population in southeastern Idaho were significantly more tolerant of the monoterpenes than those from a Montana population. For the Montana population, agar diet studies showed that camphene, myrcene, terpinolene, bornyl acetate, and tricyclene adversely affected larval growth rate and pupal weight. For the Idaho population, agar diet studies showed that only terpinolene and bornyl acetate adversely affected larval growth and pupal weights. For both populations bornyl acetate was the most toxic, and this compound in addition to other monoterpenes represent defensive mechanisms in the current year's growth of Douglas fir. Sesquiterpene and phenolic-flavonoid glycoside fractions extracted from Douglas fir current year's growth also were bioassayed using agar diets. The sesquiterpene fraction showed a significant negative effect on budworm larval growth, but phenolics and flavonoid glycosides had no effect. Sesquiterpenes, in combination with tricyclene, camphene, myrcene, limonene, terpinolene, and the acetate fraction appear to represent an effective mixture of defensive compounds against the budworm.

Monoterpene accumulation during maturation of the current year's growth of Douglas-fir, Rocky Mountain variety, is characterized by a two-phase response. Beginning at or near budbreak, α-pinene and a tricyclene-(1)-camphene-bornyl acetate biogenetic group accumulate 3–10-fold. This is followed by another lesser increase later in the season of the minor components (E-) ocimene, citronellal, and borneol. In contrast, levels of β-pinene, myrcene, and terpinolene, and the sesquiterpenes γ-cadinene, α-bisabolene, and an oxygenated sesquiterpene, did not vary significantly during needle maturation. These increases occur first in the lower canopy and proceed upward toward the mid- and upper-canopy levels shortly thereafter. Implications of these observations with respect to metabolism, host-insect interactions, and silviculture management are discussed.

We conducted a field experiment to assess interrelationships between leaf- tissue secondary chemistry, avian predation, and the abundance and diversity of arthropods occurring on sagebrush (Artemisia tridentata) in central Oregon. Arthropods were removed from individual shrubs, some of which were then caged to exclude birds. Secondary chem- istry and arthropods were sampled at intervals up to 56 wk following the defaunation/ caging treatment. Recovery rates differed among arthropod taxa and functional groups. Several sap- sucking homopterans and hemipterans reached control levels within 2-4 wk of the treat- ment, whereas abundances ofparasitoids and predators recovered to match control numbers only 6 wk after defaunation. Abundances of several herbivorous leaf-chewing taxa (pri- marily lepidopteran larvae) remained significantly depressed even 56 wk after the treatment. Fungivores (oribatid mites) reached greater abundances on defaunated than on control shrubs by the end of the experiment. There were also significant changes in the concen- trations or frequency of occurrence of several chemical compounds following the defauna- tion treatment. Several hydrocarbons, sesquiterpenes, and monoterpenes that were present in most shrubs exhibited sharply reduced concentrations in leaf tissues within 4 wk of the treatment, whereas some alcohols and ketones (linalool, borneol, thujone), which occurred at relatively low frequencies among control plants, increased dramatically in their fre- quencies of occurrence following arthropod removal. Both changes persisted for the du- ration of the experiment. We found several significant associations between the abundance or occurrence of arthropod taxa or groups and concentrations or frequencies of secondary compounds, but these were most prevalent among the leaf-chewing lepidopterans. We suggest that the shrubs responded to the removal of herbivorous, leaf-chewing arthropods by altering chemical allocation patterns; the changes in chemistry persisted for over a year because recolonization of the defaunated plants by these herbivores was slow. Effects of the caging treatment were much less obvious. The recovery of the diversity of arthropods known to constitute prey for birds in this system was slower on shrubs

The current year's growth of Douglas fir contains galactose, unusual in that this carbohydrate makes up 78.7% of the total carbohydrate fraction. An agar diet study was undertaken to determine the effects of galactose, other carbohydrates, and terpenes on western spruce budworm larval mortality, growth rate, and adult biomas production. All concentrations of the carbohydrates and terpenes tested, as well as other mineral elements not tested, were typical of the current year's foliage of Douglas fir. In experiment I, the diet containing 5.61% total carbohydrate did not significantly affect larval mortality when compared to the control diet. However, diets containing 9.45% and 15% total carbohydrate concentrations significantly increased larval mortality 64% and 96.1%, respectively, when compared to the control. Also in experiment I, terpenes alone (78.9% morality) and terpenes in combination with 9.45% and 15% total carbohydrates significantly increased larval mortality (97.2% and 100%, respectively) when compared to mortality on the control diet (44%). To determine which carbohydrate was causing the adverse effect, 6% glucose, 6% fructose, and 6% galactose were placed individually and in combination with terpenes in diets in experiment II. The 6% galactose diet significantly increased larval mortality and reduced growth rate when compared to the control, glucose, and fructose diets. Glucose resulted in 16% less larval mortality, significantly enhanced female larval growth rate and pupal weight, but did not affect male larval growth rate and pupal weight, when compared to the control. Fructose resulted in a significant decrease in larval mortality and a general trend of enhanced female and male larval growth rate and pupal weight. Larval mortality on terpenes alone was not significantly different from the control, but terpenes with 6% galactose increased larval mortality and decreased female and male growth rate and pupal weight significantly when compared to glucose-terpene and fructose-terpene diets. No significant interactions were found between carbohydrates and terpenes in either experiment.

The correlation between total phenolics and tannin content within a species is often considered to be suitably strong to allow researchers to assume, with some degree of confidence, that levels of one will approximately parallel the other. However, the manipulation of resource availability could lead to disproportionate changes in total phenolics and tannins and/or in the specific monomers of which these fractions are composed, thus altering the correlation between these components. In order to test this hypothesis, we examined the correlation between foliar levels of total phenolics (as measured by the ferric chloride assay) and tannins (as measured by an astringency assay) in Douglas-fir (Pseudotsuga menziesii Mirb. Franco) before and after the manipulation of nitrogen and water availability. Prior to manipulation of resources, the correlation between total phenolics and tannins was strong and highly significant (r2=0.869;P < 0.001). This correlation was considerably weaker and not statistically significant following resource manipulation (r2 = 0.392; 0.20

An agar diet study using western spruce budworm populations from Idaho and New Mexico was carried out to determine the effects of varying concentrations of nitrogen, beta-pinene, and bornyl acetate on larval growth and survival. Increased availability of nitrogen resulted in increased larval growth rate and survival to the adult stage. Larval growth rates from the Idaho population were higher on the high-nitrogen diet than were growth rates from the New Mexico population. The high level of beta-pinene improved larval growth at the high-nitrogen concentration. Bornyl acetate significantly reduced larval growth at both the low and high levels of nitrogen although the effect was greatest with the high-nitrogen diet. High bornyl acetate concentrations also significantly reduced survival to the adult stage. At high-nitrogen levels, a high concentration of bornyl acetate reduced larval growth rates and adult survival to a level similar to that occurring at the low nitrogen and low bornyl acetate concentrations.

Measurements of foliage quality, physiological, and phenological condition of sample trees were used as independent variables in multiple correlation analyses to determine their effect on female and male spruce budworm larval dry weights. Female budworm from trees high in foliar concentrations of beta-pinene, myrcene and total nitrogen weighed less than those from trees lacking these characteristics. Male budworm from trees high in foliar concentrations of alpha-pinene, myrcene, terpinolene, citronellyl acetate, and bornyl acetate weighted less than those from trees lacking these characteristics. Additionally, relatively vigorous and productive trees tended to be less susceptible (as evidenced by reduced larval weight) to budworm of either sex.

Data collected on natural populations of Asarum caudatum in western Washington indicated that populations were polymorphic for growth rate, seed production, and palatability to a native slug. However, in habitats where slugs were not as abundant, populations of wild ginger were dominated by individuals allocating more energy to growth rate and seed production, and less to the production of an antiherbivore mechanism. On the basis of these data, I postulated that in the absence of grazing pressure, palatable, faster growing individuals would be at a competitive advantage over slower growing individuals that devote more energy to an antiherbivore mechanism. Under grazing pressure, however, fitness of the unpalatable individuals would be increased since smaller amounts of photosynthetic tissue would be consumed. See full-text article at JSTOR

Theoretical considerations and evidence from the literature suggest that early successional plant species should make a lesser commitment of resources to defense against herbivores, and should then provide better food sources for generalized herbivores than later successional and climax plants. Commitment to defense by plants is estimated by determining short-term palatabilities to two slug species; one native to western Washington, Ariolimax columbianus, and one introduced from Europe, Arion ater. Results of tests with 100 plant species of three growth forms and from different seral stages indicated that early successional annuals were significantly more palatable than early successional perennials which were sig- nificantly more palatable than later successional species. No correlation was found between palatability and evolutionary association of the herbivores with the plant species.

The diversity of chemical identity and mode of action of toxic substances present in plants is suggested to be partially the result of generalist herbivore grazing pressure, while convergence in digestibility-reducing substances and their method of function is due primarily to specialist herbivore. Analysis of alkaloid patterns from herbaceous and woody perennials show that most species differ in their alkaloid composition and many sympatric species and subspecies show unique patterns. The high degree of functional convergence in digestibility-reducing substances is supported by convergent patterns in tannins among woody perennials.

Host plant preferences for 34 insect herbivore species are reported. Most polyphagous herbivores feeding on annuals, herbaceous perennials, and woody perennials show distinct preferences for the least abundant plant species among their various host plants. In addition, some populations of widely distributed polyphagous species are much more specialized in their diet than host plant lists alone would suggest. The high level of polyphagy on annuals and herbaceous perennials is suggested to be strongly influenced by the unpredictability of the host plant that is, in turn, controlled by environmental variability. Oligophagous herbivores preferred the least abundant woody perennials on the study sites. Ten of the 22 monophagous herbivores preferred the rarest of all the plant species on the same sites.

Litter decomposition in terrestrial habitats is affected by many factors, including temperature, moisture, and nutrient and organic composition of litter. Among organic components, lignin is the primary controlling factor of decomposition rates of surface litter during the later phase of decomposition in most habitats and during the initial phase in warm, moist habitats (i.e., those with a high actual evapotranspiration, AET). In habitats with moderate AET's, we suggest that the decreased control by lignin over annual decomposition rates of surface litter is due, at least in part, to a significant periodic or seasonal influence of other carbonbased plant secondary metabolites over rates in the initial phase of decomposition. The influence of other secondary metabolites over decomposition rates should be a function of other correlates of AET: the phytochemical composition of the community and the persistence of various secondary metabolites in litter. As AET decreases from the highest extreme, we expect mo...

Leaf tissue preferences of monophagous, oligophagous, and polyphagous insect herbivores were determined using young and mature leaf tissue abundances and herbivore feeding observations. Larvae of monophagous and oligophagous herbivores preferred young leaf tissues while, overall, larvae of polyphagous species preferred mature leaves of their various host plants. Even though a species is often polyphagous over its geographical range, larvae from local populations may be very specialized in their diet. When this occurs these specialized larvae prefer the more nutritious and perhaps more toxic young leaves of some of their host plants. Resource abundance and plant chemistry are discussed as major factors influencing herbivore feeding patterns.

Forty plant species common in western Washington were investigated for the presence of allelopathic chemicals capable of influencing the distribution of subordinate species. Under laboratory conditions most species contained inhibitory volatile compounds and many contained effective water—soluble compounds. The results of up to 13 bioassays per species were compared with descriptions of the distribution of subordinate species under natural conditions. In several cases, notably Acer circinatum, Arbutus menziesii, and Rhododendron albiflorum, high inhibition values in laboratory experiments were associated with distinct changes in ground—cover composition beneath the canopy of these species. The ratio between inhibition values produced by volatile and water—soluble inhibitors was higher in species from arid regions than in those from humid regions. This finding supports Muller and Whittaker's hypothesis that allelopathic compounds are primarily waste products.

Tannin, cell wall, and nitrogen composition of green foliage and needle litter of similar-aged Douglas-fir (Pseudotsuga menziesii Mirb. Franco) from two stands differing in density and crown closure were compared. Trees in the closed-canopy stand had a lower basal area growth rate than those in the open-canopy stand. Stands did not differ in wood basal area/ha or forest floor C/N ratios, but the closed-canopy stand had a significantly larger accumulation of forest floor biomass and significantly higher levels of field-extractable nitrogen and nitrogen mineralization rates. Green foliage from trees in the closed-canopy stand had significantly lower nitrogen, astringency, and lignin contents, but higher cellulose concentration than trees in the open-canopy stand. These trends, inconsistent with the inverse relationship often observed between nitrogen and polyphenol contents of foliage, may result from differences in relative resource availability in the two stands. In contrast to green foliage, needle litter from the two stands had comparable contents of nitrogen, cellulose, and lignin, but astringency was significantly higher in litter from the closed-canopy stand. It is suggested that, within the constraints imposed by site conditions, evergreens may alter the tannin composition of senescing foliage, potentially affecting herbivory and decomposition differently.