Apparency of pulsed and continuous pheromone to male gypsy moths

Quiescent male gypsy moths (Lymantria dispar) exposed in a wind tunnel to either pulsed (0.5-, 2- or 5-sec on, followed by a two-fold time interval off) or continuous streams of synthetic pheromone responded similarly in the proportions and latencies of wing fanning. Similarly, upwind anemotactic flight tracks in pulsed (1-sec on and 1-sec off) and continuous plumes of pheromone were indistinguishable. These data suggest that in the gypsy moth (1) pulsed pheromone stimuli would not lower the threshold, despite the improvement in the signal-to-noise ratio; and (2) temporal modulation of the pheromone plume at 1-sec intervals does not alter the preprogrammed upwind flight pattern.Journal paper No. 10916 of the Michigan State University Agricultural Experiment Station.     

Mate location strategies of gypsy moths in dense populations

In high-density populations, the male gypsy moth (Lymantria dispar) appetitive flight is primarily vertical and within 1–50 cm of tree boles. Preceding location of a female, males land on trees or occasionally tree foliage and walk while wing-fanning. In high-density populations females may mate before calling, often prior to wing expansion or hardening. Additional matings may occur before any or full deposition of egg masses. Virgin females are not coy, whether their wings are unexpanded, or hardened and held rooflike, or whether they are calling or not. They generally mate with the first willing male and do not exercise sexual selection by an acceptance or rejection reaction. The mating structure in dense populations would seem to reduce selective pressure for female emission of and direct male anemotactic flight to attractant pheromone. A low proportion of males search appropriate objects, especially tree trunks, in the walking-wing-fanning state. Such males often are successful in locating virgin females before calling commences. This male strategy presumably would not be successful in low population densities.Michigan Agricultural Experiment Station Journal Article No. 10253.     

Detoxication activity in the gypsy moth: Effects of host CO2 and NO 3 − availability

We investigated the effects of host species and resource (carbon dioxide, nitrate) availability on activity of detoxication enzymes in the gypsy moth,Lymantria dispar. Larvae were fed foliage from quaking aspen or sugar maple grown under ambient or elevated atmospheric CO₂, with low or high soil NO ₃ ^– availability. Enzyme solutions were prepared from larval midguts and assayed for activity of cytochrome P-450 monooxygenase, esterase, glutathione transferase, and carbonyl reductase enzymes. Activity of each enzyme system was influenced by larval host species, CO₂ or NO ₃ ^– availability, or an interaction of factors. Activity of all but glutathione transferases was highest in larvae reared on aspen. Elevated atmospheric CO₂ promoted all but transferase activity in larvae reared on aspen, but had little if any impact on enzyme activities of larvae reared on maple. High NO ₃ ^– availability enhanced activity of most enzyme systems in gypsy moths fed high CO₂ foliage, but the effect was less consistent for insects fed ambient CO₂ foliage. This research shows that gypsy moths respond biochemically not only to interspecific differences in host chemistry, but also to resource-mediated, intraspecific changes in host chemistry. Such responses are likely to be important for the dynamics of plantinsect interactions as they occur now and as they will be altered by global atmospheric changes in the future.     

Effects of nitrogen and Douglas-fir allelochemicals on development of the gypsy moth,Lymantria dispar

Two experiments were conducted to examine the influence of foliar nitrogen, terpenes, and phenolics of Douglas-fir on the development of gypsy moth larvae. In the first experiment, foliar concentrations of nitrogen and allelochemicals were manipulated by fertilizing 3-year-old potted seedlings with 0 or 200 ppm nitrogen. Concentrations of foliar nitrogen (0.33–2.38%) were negatively correlated with the phenolics (15.8–24.4 mg/g). Sixth-instar larvae previously reared on current-year Douglas-fir needles were allowed to feed on these seedlings. Pupal weights (312.8–995.6 mg) were positively correlated with levels of foliar nitrogen, negatively correlated with amounts of foliar phenolics, and uncorrelated with terpene concentrations. In the second experiment, terpene and phenolic extracts from Douglas-fir foliage were incorporated at natural levels into artificial diets with high and low levels of protein nitrogen. Neonate larvae grew faster and were larger on the high nitrogen control diet (4.1–4.5%), however, fourth instars performed better on the control diet with low nitrogen levels (2.5–2.7%). Foliar terpenes incorporated into diet had little effect on neonate fitness, but may induce subtle physiological changes in later instar larvae. Phenolics, alone or in combination with terpenes, excessively suppressed growth and survival, with no individuals living through the fourth instar, regardless of the nitrogen level. Incorporating foliar phenolic extracts into artificial diet caused unnatural levels of toxicity and failed to clarify the effects of Douglas-fir phenolics on gypsy moth fitness. Foliar nitrogen is a key factor influencing gypsy moth development on Douglas fir, but may be mitigated to some degree by phenolics.     

Chemical ecology of the luna moth

The effects of food plant on larval performance and midgut detoxification enzymes were investigated in larvae of the luna moth,Actias luna. Neonate larvae were fed leaves of black cherry, cottonwood, quaking aspen, white willow, red oak, white oak, tulip tree, paper birch, black walnut, butternut, or shagbark hickory. First instar survival, larval duration, and pupal weights were monitored as indices of food quality. Midgut enzyme preparations from fifth instars were assayed for -glucosidase, quinone reductase, polysubstrate monooxygenase, esterase, and glutathione transferase activities. Larval survival on seven of the 11 plant species, including several recorded host plants, was extremely poor. Larvae performed well, and quite similarly, on birch, walnut, butternut, and hickory. Activities of all enzyme systems except -glucosidase were significantly influenced by larval host plant. Of the systems assayed, quinone reductase and glutathione transferase activities were especially high. Comparisons of these values with published values for other Lepidoptera support the hypothesis that these enzyme systems are involved in conferring tolerance to juglone and related quinones occurring in members of the plant family Juglandaceae. Results suggest that host plant utilization by luna is more specialized at the individual or population level than at the species level and that biochemical detoxification systems may play a role in such specialization.     

Orientation behavior and effect of experience and laboratory rearing on responses ofCotesia melanoscela (Ratzeburg) (Hymenoptera: Braconidae) to gypsy moth silk kairomone

Cotesia melanoscela responds the same way to the silk of its host, gypsy moth larvae,Lymantria dispar L., whether or not it has previously been exposed to hosts. Parasites maintained in the laboratory for six or 25 generations also responded similarly, suggesting that orientation behavior is not modified by laboratory rearing. Details of silk-related orientation behavior were analyzed by tracing paths of female parasites when on or off silk-covered substrates. Females made 31 ° average turns every 0.5 cm whether or not they were examining silk. They tended to turn back onto a silked area when encountering a border, and their speed of movement when not examining silk was about three times higher than when examining silk.     

Pheromone-mediated behavior of the gypsy moth

The pheromone-mediated behavior of gypsy moth males was studied in both natural and simulated populations in central Pennsylvania. Feral males released into 50-m-diam plots, each with 2 feral females around the perimeter, oriented initially to trees and not to females. Neither exposure to virgin females nor exposure to wicks baited with approx 6 mg disparlure affected the subsequent sexual activity of males released into the 0.2-hectare plots. Males released into untreated plots, following 24 hr exposure in an area treated with approx 37 g/hectare of microencapsulated disparlure, located and mated with feral females within 4 min after release. None of the released males was caught in disparlure-baited Delta traps. In the disparlure-treated plot none of the females was mated. Males within this treated plot continued to search actively but did not settle down on the bark surface and initiate short-range (< 15 cm) search behavior. In plots testing the effect of various ratios of baited wicks to virgin females on disruption, there was no evidence of mating disruption due to point-source confusion. There were no significant differences in the responses of feral males to either virgin females or the various portions of Hercon wicks placed out in 0.2-hectare plots. In a series of tests using feral virgin females given various treatments to alter their physical and chemical characteristics (i.e., removed wings, denuded abdomen, washed in xylene, etc.), all females elicited the full range of sexual behavior responses of the male moths in natural populations. Apparently, males stimulated by pheromone are capable of using a number of different additional stimuli to initiate and terminate short-range sexual behavior patterns. Lymantria dispar (L.); Lymantriidae: Lepidoptera.Authorized for publication on April 19, 1976 as paper no. 5066 in the journal series of the Pennsylvania Agricultural Experiment Station, Experiment Station Project No. 2044.     

Leaf phenolic inhibition of gypsy moth nuclear polyhedrosis virus Role of polyhedral inclusion body aggregation

Bioassays with nuclear polyhedrosis virus (NPV) administered to gypsy moth larvae on leaf disks from various tree species reveal strong viral inhibition by some tree species. Phenolic extracts from inhibitory tree leaves cause virus polyhedral inclusion bodies (PIBs) to form large aggregations. However, aggregated PIBs treated with leaf extracts and administered to larvae on laboratory diet (without phenolics) retain virulence. Our results suggest that leaf phenolics, especially hydrolyzable tannins, inhibit NPV infection, but may not act via aggregations formed in the foregut.     

Capture of gypsy moth,Lymantria dispar (L.), andLymantria mathura (L.) males in traps baited with disparlure enantiomers and olefin precursor in the People's Republic of China

Pheromone traps baited with (+)-disparlure,cis-7,8-epoxy-2methyocta-decane, captured males ofLymantria dispar, L. monacha, andL. mathura in northeastern People's Republic of China.L. dispar responded to the addition of olefin to (+)-disparlure-baited traps in a negative doseresponse manner. Observations on site and seasonal capture ofL. dispar andL. mathura are discussed.Lepidoptera: Lymantriidae.     

Variation in attraction of individual male gypsy moths to (+)- and (±)-disparlure

Gyspy moth (Lymantria dispar L.) males atttracted to either (+)-cis-7,8-epoxy-2-methyloctadecane [(+)-disparlure] or its racemate [(±)-disparlure] do not represent distinct behavioral classes of individuals. Males attracted to within 2 cm of either (+)- or (+-)-disparlure dispensers on a tree trunk were caught, marked as to the lure visited, and then released into a 5×5 grid at ca. 20-m spacing of traps baited alternately with (+)-or (±)-disparlure. Males then reattracted to these lures showed no evident preference for the lures at which they were first captured, indicating that males attracted to either of these lures are not behavioral phenotypes.Michigan Agricultural Experiment Station Journal Article No. 8895.     

Gypsy moth (Lymantria dispar L.) attraction to disparlure enantiomers and the olefin precursor in the People's Republic of China

Pheromone traps baited with disparlure,cis-7,8-epoxy-2-methyl-octadecane, captured males ofLymantria dispar, the gypsy moth, at two widely separated locations in the People's Republic of China. The (+) enantiomer of disparlure attracted significantly more males than the racemate; addition of olefin reduced captures. The duration of the flight period was longer (eight weeks) and peaked earlier near Beijing than farther north near Dunhua (five weeks).Lepidoptera: Lymantriidae.     

White alder and Douglas-fir foliage quality and interegg-mass influences on larval development of gypsy moth,Lymantria dispar

Individual families of gypsy moth collected from a single population exhibited different degrees of fitness when fed diets of white alder, a suitable broadleaf host, and Douglas-fir, an unsuitable conifer host. Members of families on diets of Douglas-fir had significantly lower survival, longer larval periods, lower pupal weights, and shorter pupal periods than members of the same families fed alder. Foliar nutritional quality, including nitrogen level and allelochemical composition (terpenes and phenols), was considered the key factor responsible for these differences. Growth parameters differed significantly for families within diet treatments, indicating that the genetic resources of a family did affect performance somewhat. The influence of a family's genetic resources on larval survival was most notable when larvae were under the greatest nutritional stress.     

Gypsy moth mating disruption: Dosage effects

Small (1-hectare) plots in a dense gypsy moth (Lymantria dispar) infestation were treated with 5, 50, or 500 g racemic disparlure, and effects on male trap catch and mating behavior were studied. Capture of males in traps baited with 1, 10, 100, or 1000 g (+)-disparlure declined as disruptant dosages increased. Traps with high levels of attractant caught moths when capture in those with lower baitings was dramatically reduced. While all disruptant dosages reduced trap catch, it was reducd at least 95% at all attractant levels at the 500-g disruptant application rate. Visual estimates indicated that male moth density was similar in treated and control plots; female mating success was reduced 6.5, 34.5, and 84% in plots with 5, 50, and 500 g/ hectare disruptant, respectively. The duration of precopulatory and copulatory periods was similar for all females that were observed mating, regardless of disruptant treatment. It is proposed that reduced trap catch and female mating success are due to effects of atmospheric synthetic disparlure (disruptant) camouflaging natural attractant point sources.Lepidoptera: Lymantriidae.     

Some chemical bases for gypsy moth,Lymantria dispar,larval rejection of green ash,Fraxinus pennsylvanica,foliage as food

Green ash is one of the few tree species rejected as food by larvae of the generalist gypsy moth,Lymantria dispar L. (Lepidoptera: Lymantriidae). Such rejection is based especially on chemicals present in green ash foliage. The gypsy moth larval feeding-inhibitory activity is contained in the ethyl acetate extractables of green ash foliage. Three representative columnchromatographed fractions of the extractables contained antifeedant activity. Individual fractions showed weaker antifeedant activity compared to the total ethyl acetate extractables. Acid hydrolysis of the extractables destroyed antifeedant activity and yielded feeding stimulant activity at higher concentrations. The aqueous extractables were not feeding-inhibitory. Compounds in the green ash extractables were separated by TLC, HPLC, CC, and GC.     

Evaluation of time-average dispersion models for estimating pheromone concentration in a deciduous forest

The Sutton and more recent Gaussian plume models of atmospheric dispersion were used to estimate downwind concentrations of pheromone in a deciduous forest. Wind measurements from two bivane anemometers were recorded every 12 sec and the pheromone was emitted from a point source 1.6 m above ground level at known rates. The wingfanning response of individually caged male gypsy moths (Lymantria dispar) at 15 sites situated 20 to 80 m downwind was used to monitor when pheromone levels were above threshold over a 15-min interval. Predicted concentrations from these Gaussian-type models at locations where wing fanning occurred were often several orders of magnitude below the known behavioral thresholds determined from wind tunnel tests. Probit analyses of dose-response relationships with these models showed no relationship between predicted dose and actual response. The disparity between the predictions of concentration from these models and the actual response patterns of the male gypsy moth in the field was not unexpected. These time-average models predict concentrations for a fixed position over 3-min or longer intervals, based upon the dispersion coefficients. Thus the models estimate pheromone concentrations for time intervals appreciably longer than required for behavioral response.Michigan Agricultural Experiment Station Journal Article No. 10914.     

Host selection byBlepharipa pratensis (Meigen), a tachinid parasite of the gypsy moth,Lymantria dispar L.

The host selection process ofBlepharipa pratensis (Meigen), a tachinid parasite of the gypsy moth,Lymantria dispar L., was investigated. Once in the host's habitat, and following contact with a recently damaged leaf edge (cut, torn, eaten), the fly orients perpendicular to the edge and moves back and forth with the front tarsi grasping the damaged edge. Oviposturing (oviposition intention) may occur. Leaf exudates appear to arrest the fly on the leaf and increase tarsal examination (searching). If an edge of a gypsy moth-eaten leaf is contacted, oviposition usually occurs. Significantly more eggs are laid when host-browsed foliage is encountered, compared to mechanically cut or damaged foliage, indicating response to a cue left by the host during feeding. The number of host-damaged leaf clusters in an area significantly enhances oviposition there; in field-cage tests, significantly more eggs (7911) were laid in simulated-crown areas with all clusters browsed, compared to the adjacent areas containing 1/2 browsed (4200 eggs) and undamaged clusters (2209 eggs). A host selection sequence is suggested and discussed.     

Reassessment of interaction between gut detergents and tannins in lepidoptera and significance for gypsy moth larvae

Surface-active lysophospholipids are thought to interfere with tannin-protein interactions and may comprise an adaptation in Lepidoptera to tannin-rich diets. We found several lysophospholipids and phospholipids at about eight times the critical micelle concentration (CMC) in the midguts of gypsy moth larvae, which exhibited appropriately reduced surface tensions. We confirmed the interfering activity of lysophosphatidylcholine (lysolecithin) in a model astringency assay using tannic acid and hemoglobin (hemanalysis), but discovered that tannic acid and red oak tannin extracts precipitate the lipid from simulated midgut solutions quantitatively, even at pH 10. Leaf tannin extracts from trees on which gypsy moths grew and reproduced poorly precipitated lysolecithin more effectively than did extracts from trees on which gypsy moths performed well. Adding tannic acid to midgut fluid elevated surface tension, and about 25% of larvae feeding on oak leaves exhibited elevated midgut surface tension, suggesting a loss of surfactants. Larvae appear able to replace lost surfactants to a limited degree. An important effect of leaf tannins, and perhaps other phenolics, may be to reduce concentrations of surface-active phospholipids in the midgut and produce lipid or other dietary deficiencies in insects.     

Fall armyworm sensitivity to flavone: Limited role of constitutive and induced detoxifying enzyme activity

We used inhibition and induction of detoxifying enzymes to determine whether these enzymes allow a generalist species (Spodoptera frugiperda; fall armyworms) to cope with ingestion of the flavonoid, flavone. Flavone induces polysubstrate monooxygenases (PSMO), general esterases (GE), and glutathioneS-transferases (GST) inS. frugiperda, yet this species is affected deleteriously by low dietary concentrations of this allelochemical. First, in a series of experiments, larvae were fed artificial diets containing increasing concentrations of flavone, either alone or with known inhibitors of either PSMO, GE, or GST enzymes. In an additional treatment, flavone and inhibitors of all three enzyme systems were administered in diets simultaneously. PSMO and GE activities were reduced in vivo by their respective inhibitors, whereas that of GST was induced or unchanged. Significant synergism of flavone's growth-reducing activity occurred at the highest concentration tested (0.125% fresh mass, fm) when the PSMO inhibitor, piperonyl butoxide, or the GST inhibitor, diethyl maleate, was added to the diet, and at 0.08% fm flavone, when combined with the GE inhibitor, tri-tolyl phosphate. In many cases, however, the additive effect (i.e., reduction in growth owing to flavone alone + inhibitor alone) was greater than the synergistic effect, and no synergism occurred in the treatment with the three inhibitors combined. In the second approach, caterpillars were preexposed to a concentration of flavone (0.02% fm) that induced these enzymes ca. 1.5- to 2.5-fold, prior to switching larvae to a diet containing a higher (growth-reducing) flavone concentration (0.125% fm). The relative growth rates (RGR) of induced larvae were significantly greater (14%) than those of the uninduced larvae on the 0.125% fm flavone diet. Additionally, in two of the three experiments, relative consumption rate (RCR) was significantly greater (7–24%) in induced compared with uninduced larvae. The variable responses to inhibitor treatment and the relatively small benefit of enzyme induction suggest that these enzyme systems have minimal impact on the detoxification of flavone inS. frugiperda, even though this allelochemical induces enzyme activity and has been reported to be metabolized in vitro.     

Comparative distribution and persistence of disparlure in woodland air after aerial application of three controlled-release formulations

Disparlure, the gypsy moth sex pheromone, was aerially applied to three plots, using a different controlled-release formulation-gelatin microcapsules, flakes of plastic laminates, or hollow fibers-in each plot. Disparlure concentrations in air were measured intermittently over a 34-day period after application. Measurements were made at four heights above ground: 0.3,2,5, and 10m. Wind speed and air temperature were measured concurrently. Disparlure was emitted from the microcapsules and fibers at very high levels for the first day or two; thereafter, all three formulations produced gradually decreasing aerial concentrations. After 32–34 days, concentrations in all plots ranged from 1.5% to 15.5% of those on the first day, although most of the pheromone remained in the formulations. Results suggest that the effective life of all three formulations for mating disruption by atmospheric permeation would be near 30 days under the test conditions, with fibers likely to have the shortest life. Concentrations varied diurnally throughout the experiment, with highest levels generally between 1400 and 2200 hr each day. Concentrations were highest at the 10-m height with microcapsules and at the 0.3- and 10-m heights with flakes, but were similar at all heights with fibers. Biological implications of the results are presented.     

Effects of genotype, nutrient availability, and defoliation on aspen phytochemistry and insect performance

Genetic and environmental variability, and their interactions, influence phytochemical composition and, in turn, herbivore performance. We evaluated the independent and interactive effects of plant genotype, nutrient availability, and defoliation on the foliar chemistry of quaking aspen (Populus tremuloides) and consequences for performance of gypsy moths (Lymantria dispar). Saplings of four genotypes were grown under two conditions of nutrient availability and subjected to three levels of artificial defoliation. Concentrations of all secondary and primary metabolites evaluated responded to at least one or more of the experimental treatments. Of the secondary metabolites, phenolic glycosides were affected strongly by genotype, less so by nutrient availability, and not induced by defoliation. Condensed tannins were strongly dependent upon genotype, soil nutrient availability, and their interaction, and, in contrast to phenolic glycosides, were induced by artificial defoliation. Of the primary metabolites, foliar nitrogen was affected by genotype and soil nutrient availability. Starch concentrations were affected by genotype, nutrient availability, defoliation and interactions among these factors. Foliar water content responded to genotype, nutrient availability, and defoliation, and the effect of nutrient availability depended on genotype. Herbivore performance on these plants was strongly influenced by plant genotype and soil nutrient availability, but much less so by defoliation. Although several of the compound types (condensed tannins, starch, and water) responded to defoliation, quantitative variation in these compounds did not contribute to substantive changes in herbivore performance. Rather, the primary source of variation in insect performance was due to plant genotype (phenolic glycoside levels), while nutrient availability (foliar nitrogen levels) was of secondary importance. These results suggest that genetic variation in aspen plays a major role in determining patterns of insect performance, whereas environmental variation, such as was tested, here is of negligible importance.