Influence of epicuticular waxes from white spruce and balsam fir on feeding behavior of the eastern spruce budworm

Fourth-instar larvae of the eastern spruce bud-worm Choristoneura fumiferanaClem. (Lepidoptera: Tortricidae) were shown in two-choice feeding tests to respond differently to epicuticular waxes from different white spruce and balsam fir trees. Larvae also showed different preferences for various wax fractions obtained by separation on silicic acid.Deceased.     

Rate of pheromone release by individual spruce budworm moths

The rate and temporal pattern of pheromone emission by single and grouped female spruce budworm moths were measured by combining the trapping of pheromone on Porapak Q with a rapid luciferase bioassay developed for aldehyde pheromone. Pheromone release occurred mainly during the scotophase of a 12:12 L:D cycle in a series of bursts (up to 50 ng/hr) with considerable variability observed between insects. Analysis of over 30 individual females showed that 30% release no detectable pheromone, 60% release between 20 and 100 ng of pheromone and 10% release greater than 100 ng in a 24-hr period. Overall, a single female (1–3 days old) released an average of 60 ± 50 ng (± S.D.) of pheromone per night with a total of 260 ± 210 ng (± S.D.) being released over its life span (～7 days). Grouped females released lower quantities of pheromone. The amount of pheromone in the glands of female moths also displayed a rhythm with the levels beings higher later in the day than at the start of the photophase.     

Landscape host abundance and configuration regulate periodic outbreak behavior in spruce budworm Choristoneura fumiferana

Landscape‐level forest management has long been hypothesized to affect forest insect outbreak dynamics, but empirical evidence remains elusive. We hypothesized that the combination of increased hardwood relative to host tree species, prevalence of younger forests, and fragmentation of those forests due to forest harvesting legacies would reduce outbreak intensity, increase outbreak frequency, and decrease spatial synchrony in spruce budworm Choristoneura fumiferana outbreaks. We investigated these hypotheses using tree ring samples collected across 51 sites pooled into 16 subareas distributed across a large ecoregion spanning the international border between Ontario (Canada), and Minnesota (USA). This ecoregion contains contrasting land management zones with clear differences in forest landscape structure (i.e. forest composition and spatial configuration) while minimizing the confounding influence of climate. Cluster analyses of the 76‐yr time‐series generally grouped by subareas found within the same land management zone. Spatial nonparametric covariance analysis indicated that the highest and lowest degree of spatial synchrony of spruce budworm outbreaks were found within unmanaged wilderness and lands managed at fine spatial scales in Minnesota, respectively. Using multivariate analysis, we also found that forest composition, configuration, and climate together accounted for a total of 40% of the variance in outbreak chronologies, with a high level of shared variance between composition and configuration (13%) and between composition and climate (9%). At the scale of our study, climate on its own did not explain any of the spatial variation in outbreaks. Outbreaks were of higher frequency, lower intensity, and less spatially synchronized in more fragmented, younger forests with a lower proportion of host species, with opposing outbreak characteristics observed in regions characterised by older forests with more concentrated host species. Our study is the first quantitative evaluation of the long‐standing ‘silvicultural hypothesis’ of spruce budworm management specifically conducted at a spatio‐temporal scale for which it was intended.     

Strains of protoplasts of Entomophthora egressa in spruce budworm larvae

Protoplasts of strain 458 (S458) and strain 521 (S521) of Entomophthora egressa had LD₅₀s of 620 and 8.8 cells/insect, respectively, for sixth-instar spruce budworm larvae. Both protoplast strains exhibited biphasic growth profiles with comparable growth rates in the larval hemocoel. The growth rates of the fungal strains increased as the total hemocyte counts declined. Hemocytopenia was greatest and most rapid in larvae containing S521 protoplasts. Protoplasts of S521 exposed to α-mannosidase and β-N-acetylglucosaminidase were as virulent as the control cells. β-Galactosidase reduced protoplast virulence.     

Taste receptor activity and feeding behaviour reveal mechanisms of white spruce natural resistance to Eastern spruce budworm Choristoneura fumiferana

The pattern of feeding of Eastern spruce budworm Choristoneura fumiferana (Clem.) (Lepidoptera, Tortricidae) is compared on foliage from white spruce Picea glauca (Moench) Voss. (Pinaceae) trees previously determined to be susceptible and resistant to defoliation by budworm. No differences are observed in electrophysiological responses from taste sensilla to aqueous extracts of the two foliage types, nor is there a preference for either extract type in a choice test. Acetone extracts from the two foliage types are both preferred to a control sucrose solution, although neither elicits a preference relative to the other. These results suggest that there is no difference in phagostimulatory power of internal leaf contents of the two foliage types. Longer‐term observation of feeding behaviour in a no‐choice situation shows no difference in meal duration, confirming the lack of difference in phagostimulatory power. However, on average, intermeal intervals are twice as long on the resistant foliage, leading to an overall lower food consumption during the assay. This result suggests an anti‐digestive or toxic effect of the resistant foliage that slows behaviour and limits food intake. Previous research has shown that waxes of the resistant foliage deter initiation of feeding by the spruce budworm and that this foliage contains higher levels of tannins and monoterpenes. The data suggest that the resistant foliage contains a post‐ingestive second line of defence against the spruce budworm.     

Sublethal effects of Bacillus thuringiensis on the spruce budworm, Choristoneura fumiferana

Exposing larvae of the spruce budworm, Choristoneura fumiferana (Clemens), to sublethal ( 50% lethal dose) levels of Bacillus thuringiensis subsp. kurstaki at various stages of their development significantly increased development time to the pupal stage and reduced pupal size and number of eggs laid per female, but did not affect the proportion of embryonated eggs. The changes in larval development time, pupal weight and fecundity depended on the larval stage that was treated. Exposure of fourth instars delayed larval development and reduced only male pupal weights with no effects on fecundity. Exposure of sixth instars delayed larval development to a lesser extent than exposure of fourth instars but had a pronounced effect on weight of both male and female pupae. The effect on pupal weight was sex dependent, as males tended to be more affected than females. The reduction in male pupal weight did not appear to influence fecundity, because the effect of exposure was explained by the change in female pupal weight. Effects on larval growth and pupal weight were proportional to the dose that was ingested during exposure, and were observed at doses as low as one-tenth of the LD₅₀. Ingestion of an LD₅₀ caused a 29 or 45% delay in development of, respectively, female or male larvae when exposed as fourth instars and a 30% reduction in female pupal weight when larvae were exposed as sixth instars.     

Disruption of male spruce budworm orientation to female moths by pheromone and pheromone analogues in a wind tunnel

Abstract. The ability of male spruce budworm moths, Choristoneura fumiferana (Clem.), to orientate to and locate virgin female moths was investigated in a laboratory wind tunnel. When presented with two caged ‘target female’ moths, more than 99% of the male moths locked on and flew directly to the females. When confined for 3 h before release, downwind of a three-dimensional array of thirty-five evenly distributed female moths (a density of 40/m³), over 90% of the males still successfully oriented to one or another of the females, although not always to the target females. With an array of thirty-five septa containing either E-11-tetradecenyl acetate on E-11-tetradecenol there was no visible change in behaviour; >90% of the males flew directly to the two target females, which indicates that neither the acetate nor the alcohol has potential for mating disruption. An array of thirty-five septa containing a 955 blend of E- and Z-11-tetradecenal, the main components of the spruce budworm sex pheromone, significantly reduced the numbers of male moths able to locate the target females, and effects were concentration-dependent. The highest concentration, which gave a release rate similar to that of a calling female, produced the same level of disruption as the thirty-five females, but sensory fatigue became a factor in addition to ‘false trail following’. Lower concentrations were significantly less effective, with no evidence of sensory fatigue. At all concentrations significant numbers of males flew to the septa (false trail following), but when male moths were left uncaged in the tunnel for a further 3h, most eventually located the females. This suggests that false trail following alone will not prevent mating. The level of disruption was not improved by the addition of 2% tetradecanal to the 955 blend. Similar levels of disruption to the 955 blend were obtained with an 80:20 blend. A 5050 blend still caused a significant amount of false trail following, but less than the other blends. The ability of male spruce budworm to orientate to off-blend trails was enhanced if the males were first exposed to the natural, female-produced pheromone.     

Water holdup capacity and residence time of red spruce and balsam fir branches

Summary This study examined the throughfall dynamics of high-elevation red spruce (Picea rubens Sarg.) and balsam fir [Abies balsamea (L.) Mill.] branches. A site was established at an elevation of 1160 m on Mt. Washington, New Hampshire, USA, and branches were collected from the canopies of mature trees. Throughfall water dynamics of branches collected in September 1988 and March 1989 were determined using a fluorescent tracer in an artificial precipitation apparatus. Water holdup capacities of spruce and fir branches from different canopy positions were similar. Spruce retained more water per unit area than fir. For rain, a three-compartment model that was used to analyze water dynamics showed that the bulk of water initially on the branch experienced very slow turnover, while intercepted water left the branch quickly. There did not appear to be any difference in throughfall dynamics between spruce and fir. Our results suggest that the initial composition of rain and mobilized dry deposition will influence the composition of water in contact with the branch for a relatively long time, as branch water composition will respond slowly to changes in precipitation composition. These predictions require field testing where sequential sampling of throughfall and precipitation occurs on a time scale equivalent to 0.1–0.2 mm of precipitation.     

Chemoreception of amino acids by female fourth- and sixth-instar larvae of the spruce budworm

An electrophysiological approach was used to record the responses of maxillary styloconic sensilla of fourth- and sixth-instar larvae of the spruce budworm Choristoneura fumiferana to 14 amino acids. One cell in the lateral styloconic sensillum was identified as an amino acid-sensitive neuron. All of the amino acids tested, except l-proline and l-arginine, were detected by this cell. Arginine did not evoke a response from either the medial or lateral styloconic sensilla. Proline evoked responses from a cell in the medial styloconic sensillum. It is known from previous behavioural work that l-proline is a phagostimulant and l-valine inhibits feeding in Choristoneura; we thus further characterized the responses to these two amino acids. For both instars, l-proline was detected as low as 0.001 mmol/l and the maximal response was at 50 mmol/l. Stimulation of fourth- and sixth-instar larvae with l-valine showed that the maximum firing frequency was obtained at 1 mmol/l. Above and below this concentration, firing frequency decreases. Sensory responses to the amino acids stimuli did not correlate with known behavioral responses to similar stimuli.     

A study of the surface proteins of Entomophthora egressa protoplasts and of larval spruce budworm hemocytes

Entomophthora egressa protoplasts either exposed to or not exposed to trypsin were not attacked by either trypsinized or non-trypsinized larval spruce budworm granulocytes. Granulocytes adhered to protoplasts exposed to papain, and this adhesion could be prevented by papainizing the hemocytes. Differences were observed in the responses of two E. egressa isolates when exposed to papain or to the papain-control solutions. Exposure of hemocytes to trypsin did not reduce either the number of Absidia repens sporangiospores per granulocyte or the percentage of granulocytes with spores, whereas, exposure to papain did. The role of surface proteins, particularly glycoproteins, in hemocyte-fungal cell interactions is briefly discussed.     

Ecological mechanisms of population change during outbreaks of the spruce budworm

Abstract.  1. Stage-specific survival and recruitment of spruce budworm were measured by frequent sampling of foliage in four outbreak populations over a 15-year period in Ontario and Quebec, Canada.
2. Patterns of change in population density during the outbreak collapse phase were closely linked to changes in survival of the late immature stages, and were determined largely by the impact of natural enemies.
3. Host-plant feedback also contributed significantly to survival patterns throughout the outbreak: annual defoliation influenced survival of fourth and fifth instars and fecundity while cumulative defoliation influenced survival of the very early larval stages (first and second) via impacts on stand condition.
4. Inclusion of this host-plant feedback reveals spruce budworm population dynamics as a function of density-related trophic interactions that vary in their order and strength of influence over time. This view re-introduces the importance of forest interactions as a component of dynamics of the spruce budworm.     

Physiology of a sucrose-sensitive cell on the galea of the eastern spruce budworm larva, Choristoneura fumiferana

ABSTRACT The lateral styloconic sensillum on the galea of the eastern spruce budworm larva Choristoneura fumiferana Clem. (Lepidoptera: Tortricidae) contains a cell which responds to sucrose. The electro-physiologial threshold for response is below 0.5 mM sucrose. The K_b for the response is 1.5 mM, and the V_max is 200 impulses/s. The physiological data are interpreted with respect to the sucrose-stimulated feeding behaviour of the larva.     

Occurrence and effects of Nosema fumiferanae infections on adult spruce budworm caught above and within the forest canopy

1 Nosema fumiferanae infections in populations of both sexes of spruce budworm Choristoneura fumiferana moths, collected live above the forest canopy (canopy moths), within the tree crown (crown moths) and in drop trays (dead moths), were examined over a 5‐year period in New Brunswick, Canada. 2 The incidence of infection and of moderate–heavy infections in canopy and crown moths of both sexes increased concomitantly with moth eclosion, indicating that N. fumiferanae retards larval/pupal development, with infected moths, particularly those having higher disease loads, emerging later in the season. 3 Infection rates differed among canopy, crown, and dead female, but not male, moths. Canopy (i.e. emigrating) females had a lower incidence of infection, lower incidence of moderate–heavy infections, and had longer forewings and higher dry weights, than crown females. These results suggest that N. fumiferanae infections negatively affect aspects of female, but not male, flight performance. Regardless of infection, forewing length and dry weight of both canopy and crown females declined over the moth flight period, but infected females in both moth types were smaller than their uninfected counterparts. Forewing lengths and dry weights of moderately–heavily infected females were most severely affected. 4 Despite high annual infection rates in parents, only a small percentage of offspring (second‐instar larvae) that established feeding sites each spring were infected, indicating that high rates of horizontal transmission occurred annually throughout the larval period. 5 The present study indicates that whether N. fumiferanae infections are a debilitating sublethal factor in spruce budworm populations depends more on the disease load than on the overall incidence of infection. The potential importance of N. fumiferanae infections on various fitness parameters related to host dispersal is discussed.