Weak responses of pine sawfly larvae to high needle flavonoid concentrations in scots pine

Responses of sawfly larvae (Hymenoptera, Diprionidae) to the flavonoid taxifolin glucoside in their host plant were studied in a laboratory experiment. Larvae ofNeodiprion sertifer andDiprion pini were raised from egg hatch to cocoon spinning on two Scots pine (Pinus sylvestris) chemotypes, one without needle taxifolin glucoside (–) and the other containing 2–4% taxifolin glucoside (+). The (+) chemotype had somewhat lower concentrations of needle terpenoids (resin acids) than the (–) chemotype. Current-year needles had higher taxifolin glucoside concentrations than mature needles. There were no differences in survival or body size betweenN. sertifer larvae that fed on the (+) chemotype and those that fed on the (–) chemotype. FemaleD. pini larvae raised on (+) needles developed 6% more slowly than larvae fed (–) needles. The results from this study are contrary to earlier findings showing that flavonoid glucosides have strong negative effects on insect performance. Possible explanations for the different outcomes are discussed.     

Effects of nitrogen fertilization on secondary chemistry and ectomycorrhizal state of Scots pine seedlings and on growth of grey pine aphid

Effects of nitrogen availability on secondary compounds, mycorrhizal infection, and aphid growth of 1-year-old Scots pine (Pinus sylvestris L.) seedlings were studied during one growing season. Seedlings were fertilized with nutrient solutions containing low, optimum, and two elevated (2 × and 4 × optimum) levels of NH₄NO₃. At the end of growing season foliar nitrogen concentration, needle biomass, needle length, water contents of needles, root collar diameter, and number of buds increased with enhanced nitrogen availability. Addition of nitrogen did not have effect on concentrations of monoterpenes in growing needles, but in mature needles significantly decreased concentrations of some individual and total monoterpenes were detected. In growing needles the concentrations of some individual resin acids decreased, and in mature needles concentrations of some individual and total resin acids increased with increased nitrogen fertilization. Higher numbers of resin ducts were found in mature needles with nitrogen fertilization. Nitrogen fertilization decreased total phenolic concentrations in growing and mature needles of the current year, but in needles of the previous year no significant differences occurred. Mycorrhizal infection was highest at medium (optimum and 2 × optimum) nitrogen fertilization levels. The relative growth rate (RGR) of grey pine aphid [Schizolachnus pineti (F.)] responded positively to the increase in foliar nitrogen content. However, the increase in aphid performance between optimum and the highest fertilization level was slight. This may indicate a deterring effect of resin acids on aphids. The results indicate that carbon/nutrient balance hypothesis fails to predict directly the effects of nitrogen availability on concentrations of carbon-based defensive compounds in mature foliage. Altered nitrogen supply affects allocation to secondary metabolites differently, depending on the developmental state of the plant and the biosynthesis pathway, cost of synthesis, and storage of compounds.     

Field responses of redheaded pine sawfly males to a synthetic pheromone and virgin females in Florida

Field tests using the sawfly pheromone [3,7-dimethylpentadecan-2-ol acetate with (–)-erythro configuration (2S,3S)] were conducted near Gainesville, Florida, during 1978–1981 to determine the attraction ofNeodiprion lecontei males to baited traps with respect to time of year and time of day. Greatest numbers of males were caught during May, July, and September in traps placed within a pine stand from July 1978 to July 1979. Males were only caught between 1400 and 2000 hr on 10 dates in June and 10 dates in September–October 1980, and 10 dates in June 1981, with greatest catches from 1600–1800 hr. Catches in a synthetic-baited trap and in virgin female-baited traps were similar with respect to time of day.University of Florida Agriculture Experiment Stations Journal Series 2920.Hymenoptera: Diprionidae.     

The chemical feeding ecology ofNeodiprion dubiosus schedl,N. rugifrons midd., andN. lecontei (Fitch) on jack pine (Pinus banksiana lamb.)

Two monophagous sawflies,Neodiprion dubiosus andN. rugifrons, actively avoid feeding on the current season's needles of jack pine,Pinus banksiana, while the polyphagousN. lecontei feeds indiscriminately on foliage of all age classes. Bioassay studies with needle extracts indicate thatN. dubiosus andN. rugifrons larval feeding preference is governed by a group of tricyclic diterpene resin acids. Resin acids commonly occur in needles of all ages; however, levopimaric, palustric, dehydroabietic, and neoabietic acids were at higher concentrations in the new needles.N. dubiosus was significantly deterred from feeding on 1-year-old foliage when treated with the pure resin acids, palustric and levopimaric, at 0.5 and 1.0 mg/ml MeOH.N. rugifrons larvae were deterred from feeding by pure dehydroabietic, neoabietic, and palustric acids at 1.0 mg/ml MeOH.     

Multivariate study of moose browsing in relation to phenol pattern in pine needles

Extracts of needles from Scots pine,Pinus sylvestris L., grown on fertile (A) or poor (B) soil, and from lodgepole pine,P. contorta Doug. (C), have been analyzed with HPLC and UV detection. By application of multivariate analysis, the phenol pattern was shown to vary between species and between habitats. Within groups A and B, moose food choice was highly correlated to phenol pattern. Concentrations of most of the analyzed compounds were internally correlated. Information related to pine growth rate could also be extracted from the phenolic pattern for A, B, and C. Moose food choice was not correlated to pine growth rate.     

Field response and gas-liquid chromatograph separation of optically active synthetic and natural pheromones in two sympatric diprionid sawflies,Neodiprion nanulus nanulus andNeodiprion sertifer (Hymenoptera: Diprionidae)

The male red pine sawflyNeodiprion nanulus nanulus Schedl responds mainly to the (2S,3S,7S)-3,7-dimethylpentadecan-2-yl acetate (2S,3S,7S-A) in the field. No other isomer is as effective by itself or syner-gizes 2S,3S, 7S-A in this species. ForNeodiprion sertifer (Geoffroy), however, we confirmed our earlier report that males responded significantly to a 50.003 mixture of 2S,3S,7S-A and either 2S,3R,7R-A or 2S,3R,7R/S-A. The 2S,3S,7S-A isomer was separated from 2S,3R,7R-A by capillary gas-liquid chromatographic (GLC) analysis using Carbowax 20 M and DB-5 columns. The latter column also separated 2S,3S,7S-A from 2S,3R,7S-A. Only 2S,3S,7S-A was found in the natural pheromones ofN. n. nanulus andN. sertifer by GLC analysis.     

Comparing the Variation of Needle and Wood Terpenoids in Scots Pine Provenances

We determined variation in both the concentration and composition of terpenoids in needles and wood within nine Scots pine (Pinus sylvestris) provenances. Seedlings of different provenances representing a 1200-km N–S transect from Estonia to northern Finland were cultivated in Suonenjoki nursery field, central Finland, for seven years. Growth of seedlings and the number of vertical resin ducts in wood were also determined. -Pinene and 3-carene were the major monoterpenes both in the needles and wood. The total monoterpene concentration was about five times higher in the needles than in the wood. A strong positive correlation was found between proportional quantities of several terpenes of the needles and wood, particularly for 3-carene, sabinene, and terpinolene. The needles contained both labdane-type and tricyclic resin acids, whereas the wood contained only tricyclic ones. The wood had a four times higher tricyclic resin acid concentration than the needles. The highest total monoterpene concentration in the needles and in the wood occurred in the most northern Muonio provenance and in the most southern Saaremaa provenance plants, respectively. The amount of high 3-carene genotype trees decreased among the northern provenances. The wood of the most northern Muonio provenance showed the lowest total resin acid concentration, but provenance did not affect total tricyclic resin acids in the needles. Korpilahti provenance trees from central Finland had the best growth in height. In addition, Korpilahti and Ruokolahti provenance trees showed largest radial growth of stem and smallest number of vertical resin ducts. The results suggest that especially the proportional quantity of 3-carene in the needles could be used in estimating the amount of this compound in the wood and vice versa.     

Correlation of phenolic acid content of maize to resistance toSitophilus zeamais,the maize weevil,in CIMMYT'S collections

The (E)-ferulic acid content of the grain of nine populations of land races of maize derived from CIMMYT's collections was found to be negatively correlated to susceptibility characteristics towards the maize weevilSitophilus zeamais. Correlation coefficients for six susceptibility parameters and (E)-ferulic acid content were significant and ranged from –0.58 to –0.79. A multiple regression analysis by the SAS forward procedure using the primary seed characteristics associated with susceptibility indicated that the ferulic acid content was the only significant factor in explaining variation in at least two susceptibility parameters: the Dobie index and adult preference. In 15 CIMMYT pools, correlations between four susceptibility parameters and (E)-ferulic acid content were also significant (–0.76 to –0.81). The results suggest that phenolic acid content is a leading indicator of grain resistance or susceptibility to insects and may represent a newly identified mechanism of resistance.     

Effects of Enantiomeric Blend of Verbenone on Response ofIps paraconfusus to Naturally Produced Aggregation Pheromone in the Laboratory

The aggregation pheromone produced by maleIps paraconfusus (Coleoptera: Scolytidae) tunneling in Monterey pine,Pinus radiata, logs was trapped on Porapak Q. A concentration of an extract of trapped volatiles that attracted beetles was determined in the laboratory through a concentration–response analysis of walking behavior of males and females. The interruptant effects of four concentrations of verbenone on response to a constant concentration of this naturally produced aggregation pheromone were tested with males and females. Independent of its enantiomeric composition [99.5% (S)-(–), 93.1 % (S)-(–), and 98.3% (R)-(+)], verbenone significantly reduced the percentage of females (but not males) reaching the attractant source. However, when the time required for beetles of both sexes taken together to reach the attractant source was considered, verbenone of higher enantiomeric purity had a greater effect on beetle behavior. Solutions of 99.5% (S)-(–)- and 98.3% (R)-(+)-verbenone increased the time required for beetles to reach the attractive source when compared to 93.1% (S)-(–)-verbenone. When pooled across enantiomeric blends, increasing concentrations of verbenone resulted in slower responses in beetles that reached the attractant source within 2 min. Males and females did not respond to verbenone alone.     

Drought Stress Alters the Concentration of Wood Terpenoids in Scots Pine and Norway Spruce Seedlings

Drought is known to have an impact on the resistance of conifers to various pests, for example, by affecting resin flow in trees. Little is known, however, about the quantitative and qualitative changes in resin when trees are growing in low moisture conditions. We exposed Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings to medium and severe drought stress for two growing seasons and analyzed the monoterpenes and resin acids in the main stem wood after two years of treatment. In addition to secondary chemistry, we measured the level of nutrients in the needles and the growth response of seedlings. After the first year of treatment, drought stress did not affect the growth of seedlings, but in the second year, shoot growth was retarded, especially in Scots pine. In both conifer species, severe drought increased the concentrations of several individual monoterpenes and resin acids. Total monoterpenes and resin acids were 39 and 32% higher in severe drought-treated Scots pine seedlings than in the controls, and 35 and 45% higher in Norway spruce seedlings. In Scots pine needles, the concentrations of nitrogen and phosphorus increased, while magnesium and calcium decreased compared to controls. In Norway spruce needles, nutrient concentrations were not affected. The results suggest that drought stress substantially affects both the growth of conifers and the chemical quality of the wood. We discuss the potential trade-off in growth and defense of small conifer seedlings.     

Effect of urea fertilizer application on soluble protein and free amino acid content of cotton petioles in relation to silverleaf whitefly (Bemisia argentifolii) populations

The impact of urea nitrogen fertilization on silverleaf whitefly, Bemisia argentifolii Bellows & Perring, population dynamics was examined in field-grown cotton (Gossypium hirsutum L.). Five urea nitrogen treatments were tested, consisting of soil applications of 0, 112, 168, and 224 kg nitrogen per hectare, and a combined soil–foliar application of 112:17 kg nitrogen per hectare. A positive response was observed between N application rates and the measured levels of nitrate N in petioles from mature cotton leaves. Similarly, a positive response was observed between N application rates and the numbers of adult and immature whiteflies appearing during population peaks. To determine whether these positive responses were related, we measured the levels of dietary N compounds (proteins and free amino acids) that would be available for insect nutrition in cotton petioles at the different N application rates. Sampling dates and N application treatments affected levels of soluble proteins in cotton petioles, and interactions between sampling dates and treatments were significant. Across all sampling dates, the relationship between N application rates and levels of soluble proteins was linear. Sampling dates also affected levels of total and individual free amino acids. Fertilizer treatments only affected levels of total amino acids, aspartate, asparagine, and arginine plus threonine. Levels of aspartate or asparagine and the N application rates were linearly correlated. No significant correlations were observed between levels of dietary N compounds in cotton petioles and numbers of whiteflies, either adults or immatures, on the cotton plants.     

Repellency of volatile fatty acids present in frass of larval yellow mealworms,Tenebrio molitor L. (Coleoptera: Tenebrionidae), to larval conspecifics

Frass of late-instar larvae ofTenebrio molitor L. contained 0.0889 g of butyric acid, 0.0279 g of propionic acid, and 0.0175 g of valeric acid per 100 g. Grouped larvae were strongly repelled by butyric acid at the 10^–1 M concentration. Lower concentrations of butyric acid were less repellent. Valeric acid was repellent at 10^–1 M to 10^–3 M concentrations, below which no tested concentration, including one identical to that occurring in prepared solutions of frass found to be attractive, displayed any effect. Propionic acid was repellent at the concentration in prepared solutions of frass, which were strongly attractive. Concentrations of 10^–1 M, 10^–3 M, and 10^–4 M propionic acid were also found to be repellent. The implications of the repellency of these compounds to groups of mealworm larvae are discussed, with particular reference to the interaction between these frass components and others that have already been studied.     

Effects of verbenone andtrans-verbenol on the response ofDendroctonus brevicomis to natural and synthetic attractant in the field

Verbenone andtrans-verbenol were investigated as candidate interruptants for use as tree protectants. Verbenone andtrans-verbenol, pheromones released byDendroctonus brevicomis during host colonization, reduced the trap catch ofD. brevicomis near sources of the attractant composed ofexo-brevicomm, frontalin, and myrcene. Catch reduction at some trap positions was greater at a high release rate than at a low release rate oftrans-verbenol alone and of the combination of verbenone plustrans-verbenol. Verbenone also reduced catches at traps baited with attractive bolts from trees under attack byD. brevicomis. Attempts to use verbenone to protect living trees fromD. brevicomis attack were inconclusive.Coleoptera: Scolytidae.This research was supported in part by the Forest Service, U.S. Department of Agriculture, and in part by grants from the National Science Foundation/Environmental Protection Agency (grant NSF-6B-34718/BMS75-04223), and in part by grants from the Rockefeller Foundation to the University of California. Mention of a proprietary or commercial product does not constitute recommendation or endorsement of the product by the U.S. Department of Agriculture and does not imply its approval to the exclusion of other products that also may may be suitable.     

Response of total tannins and phenolics in loblolly pine foliage exposed to ozone and acid rain

Tannin and total phenolic levels in the foliage of loblolly pine (Pinus taeda L.) were examined in order to evaluate the effect of atmospheric pollution on secondary plant metabolism. The trees were exposed to four ozone concentrations and three levels of simulated acid rain. Tannin concentration (quantity per gram) and content (quantity per fascicle) were increased in foliage exposed to high concentrations of ozone in both ozone-sensitive and ozone-tolerant families. No effect of acid rain on tannins was observed. Neither total phenolic concentration nor content was significantly affected by any treatment, indicating that the ozone-related increase in foliar tannins was due to changes in allocation within the phenolic group rather than to increases in total phenolics. The change in allocation of resources in the production of secondary metabolites may have implications in herbivore defense, as well as for the overall energy balance of the plant.AAES Journal no. 9-902690P.     

Male-specific conversion of the host plant compound,myrcene, to the pheromone, (+)-ipsdienol,in the bark beetle,Dendroctonus brevicomis

When both sexes of the bark beetle,Dendroctonus brevicomis LeConte, were exposed to vapors of myrcene from ponderosa pine, only the male produced (+)-ipsdienol. In the field, racemic ipsdienol significantly reduced the attraction of both sexes in flight to a mixture of myrcene and the aggregation pheromones,exo-brevicomin and frontalin. This suggests that ipsdienol may be involved in regulating colonization density ofD. brevicomis. The implications of the biosynthesis of various enantiomers of ipsdienol byD. brevicomis and the cohabitating bark beetles,Ips paraconfusus andI. pini, in relation to their behavioral responses are discussed in regard to reducing interspecific competition.Coleoptera: Scolytidae.