Branch-cutting behavior by the vole (Microtus pennsylvanicus)

While investigating the preferences of meadow voles (Microtus pennsylvanicus) toward various coniferous species, we noted they often cut the branches of young trees, and then left them lying on the snow for two days or more before eating them completely. We tested whether this behavioral manipulation of a potential food resource resulted in a decrease in the level of secondary metabolites in branches. We found that after being cut for more than three days, the branches of young conifers had significantly less phenolics and condensed tannins. These postmanipulation levels of secondary metabolites were similar to those found in preferred summer food resources. Voles were thus capable of transforming a low-quality resource into one that was more compatible with their feeding patterns and detoxification mechanisms.     

Preference of meadow voles (Microtus pennsylvanicus) for conifer seedlings: Chemical components and nutritional quality of bark of damaged and undamaged trees

Meadow voles (Microtus pennsylvanicus Ord.) of the 1987–1988 population density peak left several seedling plantations of Norway spruce (Picea abies) and Norway pine (Pinus resinosa) severely damaged in southern Quebec, Canada, while white spruce (Picea glauca) and white pine (Pinus strobus) suffered no damage. We compared levels of crude protein, total nonstructural carbohydrates (TNC), total phenols, and monoterpenes in the bark to detect differences between damaged and undamaged seedlings within and between plantations. There were no significant differences in levels of nutritional components between both types of seedlings within the same plantation. However, differences were noted among tree plantations for protein, TNC, and phenolics content but these differences could not be assigned to the voles' selection for particular species. Quantitative and qualitative differences also were registered in monoterpene levels between species. -Myrcene and bornyl acetate were found only in seedlings of undamaged species. Moreover, the latter contained higher levels of limonene. We conclude that the presence of specific monoterpenes is more important in defense mechanisms of conifer seedlings than relative levels of nutrients (protein or TNC) or total phenolics.     

Plant phenolics as chemical defenses: Effects of natural phenolics on survival and growth of prairie voles (Microtus ochrogaster)

Very few studies have shown experimentally that plant chemical defenses actually reduce the performance of individual mammalian herbivores, much less the density of mammalian populations. We investigated the effects of representatives of three classes of plant phenoiics on the survival and growth of prairie voles by incorporating the compounds into artificial diets and feeding them to weanlings for three weeks. At low levels of protein, both quercetin (a flavonoid) and tannic acid (a hydrolyzable tannin) caused reduced growth rates; no effect occurred at high levels of protein. Quebracho (a condensed tannin) inhibited feeding and thus was lethal at all levels of protein. These results indicate that plant phenolics are likely to influence the performance and dynamics of natural populations of microtine rodents by reducing the quality of available forage. The hypothesis that the primary mode of action of the phenoiics is the reduction of digestibility of protein was not supported. The reduced growth caused by both quercetin and tannic acid could be attributed primarily to their toxicity. The effect of quebracho resulted from reduced intake (unpalatability).     

Lespedeza phenolics andPenstemon alkaloids: Effects on digestion efficiencies and growth of voles

Lespedeza cuneata contains high levels of phenolics and is a common food plant of the meadow vole (Microtus pennsylvanicus);Penstemon digitalis contains substantial quantities of alkaloids and is a common food plant of meadow voles and prairie voles (M. ochrogaster). We investigated the palatability of these plants and the effects of their secondary compounds on the digestion efficiencies and growth of both species of voles. Voles ate very little of either plant when alternative food was present. Phenolics and alkaloids were extracted from the plants, incorporated into separate artificial diets, and fed to weanling voles for three weeks.Lespedeza phenolics reduced the growth of meadow voles but not prairie voles throughout the feeding trial. These compounds disrupted digestion, reducing protein digestibility by more than half.Penstemon alkaloids lowered only the initial growth rates of prairie voles and had no significant effect on meadow voles. Prairie voles tended to increase food consumption rates on bothLespedeza andPenstemon diets. This response offset some of the decrease in digested protein intake in the first case, and offset the increased metabolic cost of processing the diet in the latter case. Our results indicate that althoughLespedeza phenolics andPenstemon alkaloids do influence consumption and digestibility of artificial diets, they do not greatly reduce the performance (i.e., growth or survival) of voles when consumed at levels generally observed for wild voles. However, these compounds probably do place an upper limit on the amount of freshLespedeza andPenstemon that can be consumed and contribute to the generalist feeding strategy of voles.     

Effect of birch (Betula pendula) bark and food protein level on root voles (Microtus oeconomus): II. detoxification capacity

The effect of protein and birch bark powder (BBP) content of forage on detoxification capacity of root voles (Microtus oeconomus) was studied. Young voles were fed with eight different diets for two weeks. Individuals on low (3%) protein diets had significantly lighter livers and kidneys than those on moderate (6%) or high (12%) protein diets. Birch bark powder addition did not have significant effect on organ weight. Detoxification was significantly induced, apparently due to secondary compounds in BBP. The activity of ethoxyresorufin-O-dealkylase (EROD) was high when protein content or BBP concentration in forage was high as compared to low protein diets or diets containing no BBP. Glucuronidation, on the other hand, was not induced by BBP. High BBP content caused serious physiological stress to the voles. The only individuals surviving were those capable of sufficiently allocating energy and protein to detoxification.     

Effect of birch (Betula pendula) bark and food protein level on root voles (Microtus oeconomus): I. food consumption,growth, and mortality

The effect of protein and birch bark powder (BBP) content of forage on food consumption and growth of root voles (Microtus oeconomus) was studied by feeding young voles with eight different diets for two weeks. Voles ate more when the protein content of forage was moderate (6%) or low (3%), and when BBP was added to the diets, as compared to a high (12%) protein diet without BBP addition. On high and moderate protein diets voles gained weight, whereas on low protein diets they lost weight. Addition of BBP caused reduced growth or weight loss. Consuming a diet with moderate (6%) protein and high BBP (20%) resulted in high mortality (9 of 10 died). These results show that root voles need at least 6% protein for growth and that the effect of birch bark depends on the protein content of the forage.     

Variation of total phenolic content and individual low-molecular-weight phenolics in foliage of mountain birch trees (Betula pubescens ssp.tortuosa)

We studied seasonal and between-tree variation in the composition and content of total and individual low-molecular-weight phenolics (LMWP) in leaves of mountain birch trees (Betula pubescens ssp.tortuosa). The major phenolic compounds were chlorogenic acid, quercetin-3-O--D-glucuronopyranoside, myricetin-3-O-(5-acetyl)-L-rhamnopyranoside, and 1-O-galloyl--D-(2-O-acetyl)-glucopyranose. The content of total phenolics, as well as the sum of individual LMWP, varied only slightly among trees while variation in contents of individual LMWP was large. Concentrations of almost all phenolics decreased during the growing season but pairwise correlations between individual phenolics remained similar over the whole season indicating tree-specific LMWP profiles over the season. Among flavonoids, the between-tree component of variation was 2.6 times as large as the seasonal component, while for variation of nonflavonoids the between-tree component was larger than the seasonal one. To explain the significant correlations within both flavonoid and nonflavonoid compounds, we discuss the biogenesis of LMWP in birch leaves, as well as their ecological role.     

Orientation behavior of second-instar larvae of eastern spruce budwormChoristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae) in a y-type olfactometer

A two-choice, wind-tunnel olfactometer was designed and contructed to determine whether second-instar eastern spruce budworm larvae,Christoneura fumiferana (Clem.), could detect and discriminate among host-plant volatiles. Volatiles of current year's growth ofPicea glauca were preferred over those ofP. rubens, P. mariana, or air.Abies balsamea was preferred overP. mariana or air.P. rubens andP. mariana were both preferred over air. Two-year-old growth ofP. glauca, A. balsamea, P. rubens, andP. mariana were all preferred over air. Current year's growth of these host evergreen species was usually preferred over former year's growth of the same species in each case.     

Comparison of release devices for stoat (Mustela ermined) semiochemicals used as montane vole (Microtus montanus) repellents

Fenced enclosures were used to simulate peak populations of montane voles (Microtus montanus) for field bioassays of a stoat (Mustela erminea) scent mixture in various controlled-release devices. A 1 1 mixture of 2-propylthietane and 3-propyl-1,2-dithiolane was dispensed in capillary tubes, clay pellets (activated alumina), rubber septa, and plastic rope. Release devices were placed near, or attached to, young apple trees planted in blocks in two enclosures containing high populations of voles. The stoat scent mixture in rubber septa and plastic rope significantly reduced vole attack of trees compared with a control, pellets, and capillary tubes. In terms of complete girdling (tree mortality), this odor in septa and rope also significantly reduced feeding compared with the control, and the rubber septa formulation was more effective than either the pellets or capillary tubes. Fenced populations of voles may be used effectively to conduct field bioassays during low years in vole cycles. Variations of release devices (with this stoat scent mixture) based on rubber septa for broadcast application to depress vole population density and survival, and plastic rope attached to individual trees to generate an avoidance response, should be used for forest and agricultural crop protection.     

Importance of lichen secondary products in food choice of two oribatid mites (Acari) in an alpine meadow ecosystem

In an alpine meadow ecosystem in the Swiss National Park, the lichensCetraria islandica andCladonia symphycarpa occur in small vegetation patches dominated byCarexfirma (Cyperaceae) andSesleria coerulea (Gramineae). Laboratory food-choice experiments with two oribatid mitesFuscozetes setosus andCarabodes intermedius show that the thallus structure and secondary products ofC. islandica are repellent factors against herbivorous mites. In contrast, secondary products ofC. symphycarpa may be an attractant for the mites.     

The role of phytoecdysteroids in bracken fern,Pteridium aquilinum (L.) Kuhn as a defense against phytophagous insect attack

Analysis of green bracken fronds collected during July, August, and October, 1975, for phytoecdysteroids showed that these compounds occur in only trace amounts (0.25–0.53 g/kg fresh weight [FW]). The effect of ecdysteroids on the feeding behavior of seven species of insect showed that four species were deterred at ecdysteroid concentrations at or above 60 mg/kg FW diet; one species of insect at 6 mg/kg or above, and two species which were not affected at the higher concentrations. It was concluded that the levels of phytoecdysteroids in bracken would not deter insects from feeding on the plant. The previously published data relevant to the possible role of phytoecdysteroids as defense compounds are also discussed.     

Phenylpropenoid phenolics in sweetbay magnolia as chemical determinants of host use in saturniid silkmoths (Callosamia)

Host plant chemistry can play an important role in determining the evolution of host use patterns in herbivorous insects by influencing host selection, consumption, and assimilation of foliage. We used a comparative approach to test the hypothesis that specialist herbivores of sweetbay magnolia (Magnolia virginiana) possess adaptations that allow them to overcome chemical deterrents or toxins that prevent herbivory by unadapted herbivores. The three silkmoth species in the genusCallosamia can be collectively regarded as specialists on magnoliaceous hosts; however, only the monophagousC. securifera is able to complete development on sweetbay magnolia, its natural host. In laboratory assays with intact foliage, bothC. angulifera and the polyphagousC. promethea fed readily on sweetbay but were unable to survive past the third instar. Two neolignan compounds, magnolol and a biphenyl ether, were found to reduce neonate growth and survival of unadapted herbivore species when painted on acceptable host leaves at concentrations similar to those found in sweetbay foliage. Both compounds significantly reduced neonate growth ofC. angulifera andC. promethea but had no effect on the sweetbay specialist,C. securifera, indicating that the latter species possesses the unique ability in the genus to tolerate, metabolize, or otherwise circumvent the phytochemical defenses of this host.     

The effects of chronic tannic acid intake on prairie vole (Microtus ochrogaster) reproduction

The hypothesis was tested that the reproductive performance of voles would be reduced when fed diets containing tannins either because of increased metabolic rates, decreased intake, or decreased digestive efficiency. We fed a ration containing 4% tannic acid (TA) (dry mass basis) to 24 pair of prairie voles (Microtus ochrogaster) and compared reproductive performance (litter size, birth weights, body mass of the young until weaning, and mass changes in the dams) to that of 24 pair of prairie vole fed a control ration. We also compared the intake rates, digestive efficiency, and metabolic rates [as measured by Vo₂ consumption (cm³/hr)] of dams and young fed both rations. Voles consuming 4% TA diets produced litters of similar size and mass as did voles fed the control ration. Furthermore, the mass of the young of dams fed the tannic acid ration were similar to the young of dams consuming the control ration through day 19 postpartum. However, dams consuming the tannic acid ration lost mass throughout lactation while the control dams maintained mass. Because the Vo₂ rates of both treated and control dams and litters were similar, we conclude increased metabolic costs were not the reason for the observed mass loss but, rather, reduced digestive efficiency, reduced intake of digestible dry mass, and apparent digestible nitrogen.     

Role of host-produced stimuli and learning in host selection behavior ofCotesia (=Apanteles) marginiventris (Cresson)

The study was conducted to determine how various factors, including learning, influence the host-selection behavior of the larval parasitoid Cotesia (=Apanteles)marginiventris (Cresson). Frass, silk, and feeding damage fromSpodoptera frugiperda (J.E. Smith) larvae elicited visits and antennal examinations by females of the parasitoid. Learning played a role in the effective response to these stimuli in that previous contact experience with hosts and/or chewed leaves, exuviae, frass, or hemolymph significantly enhanced the response of the parasitoid. Previous contact with host plants alone did not improve the responses. Experienced parasitoids were more active and spent less time at rest than inexperienced parasitoids. Previous experience was also found to be of significant importance in the ability of the parasitoid to discriminate between parasitized and unparasitized host larvae.Hymenoptera: Braconidae.In cooperation with the University of Georgia College of Agriculture Experiment Stations, Coastal Plain Station, Tifton, Georgia 31793-0748.Mention of a proprietary product in this paper does not constitute endorsement of this product by the USDA.Visiting scientist with the Department of Entomology, Coastal Plain Station, Tifton, Georgia, during this study.     

Role of (Z)- and (E)-11-tetradecenyl acetate pheromone components in the sexual behavior of the (Z) strain of the european corn borer,Ostrinia nubilalis (Lepidoptera: Pyralidae)

A glass tube olfactometer bioassay was used to examine pheromone response of males of the (Z)-pheromone strain ofOstrinia nubilalis (Hubner). The presence of (E)-11-tetradecenyl acetate at the natural ratio to (Z)-11-tetradecenyl acetate (973; ZE) did not consistently elevate wing-fanning, upwind walking, or clasper extrusion over (Z)-11-tetradecenyl acetate alone. This bioassay did not reveal the behavioral role of (E)-11-tetradecenyl acetate.Published as Journal Article No. 10264 of the Michigan State University Agricultural Experiment Station.     

Role of volatile inforchemicals emitted by feces of larvae in host-searching behavior of parasitoidCotesia rubecula (Hymenoptera: Braconidae): A behavioral and chemical study

The role of volatile infochemicals emitted by feces of larvae in the host-searching behavior of the parasitoidCotesia rubecula was evaluated during single- and dual-choice tests inside a wind tunnel. The following treatments were tested: feces produced by second and fourth instars ofPieris rapae (preferred host), second instars ofP. brassicae (inferior host), second instars ofP. napi (nonhost), and wet feces of second instars ofP. rapae. During a single-choice situation females ofC. rubecula oriented to all types of feces tested. When a preference was to be made,C. rubecula preferred feces of second instars ofP. rapae over that of fourth, feces ofP. rapae over that ofP. brassicae, feces ofP. napi over that ofP. brassicae, and wet over normal host feces. No preference was exhibited between feces of second instars ofP. napi and that of second instars ofP. rapae. The relative importance of infochemicals from host feces versus plant damage caused by host larvae to the searching behavior ofC. rubecula was also evaluated. Plant damage was more important to the searching females than host feces when feces were present in specific concentrations in relation to damage. The volatiles released by normal and wet feces of second instars ofP. rapae, wet feces of fourth instars ofP. rapae, and normal and wet feces ofP. brassicae were collected and identified. Overall, 85 chemical compounds were recorded belonging to the following chemical groups: alcohols, ketones, aldehydes, esters, isothiocyanates, sulfides, nitriles, furanoids, terpenoids and pyridines. The blend of chemicals emitted by feces of different instars ofP. rapae and different species ofPieris exhibited an instar and species specificity in both quantity and quality. Wetting of normal feces increased the amount of volatile chemicals released, and it was also responsible for the appearance of new compounds. The role of feces of larvae in the host-seeking behavior ofC. rubecula is discussed.     

Role of chirality in olfactory-directed behavior: Aggregation of pine engraver beetles in the genusIps (Coleoptera: Scolytidae)

There has been a renaissance of interest in the significance of enantiomeric composition in biological systems. Three chiral monoterpene alcohol aggregation pheromone components (ipsenol, ipsdienol, andcis-verbenol) commonly isolated from engraver beetles (Ips spp.) provide a paradigm for this theme as it relates to olfactory-guided insect behavior. The literature pertaining to this system is reviewed and the effects of the enantiomeric composition of these semiochemicals on theIps spp. community is explored on two trophic levels. Hypotheses generated from the well-studied aggregation pheromone production and response patterns forI. paraconfusus Lanier andI. pini (Say) are generalized to the North American species in the genus. Despite the progress withI. paraconfusus andI. pini, substantial deficiencies exist in our understanding of the role of enantiomeric composition in pheromonal/allomonal effects in different subgeneric groups, in the regulation and mechanisms of stereoselective biosynthesis of the monoterpene alcohols, and in the benefits derived by individual insects that produce relatively large proportions of inactive or interruptive enantiomers with attractive enantiomers.     

Defensive mechanisms of loblolly and shortleaf pine against attack by southern pine beetle,Dendroctonus frontalis Zimmermann,and its fungal associate,Ceratocystis minor (Hedgecock) Hunt

Loblolly and shortleaf pine growing on a single site in the North Carolina piedmont were examined to determine similarities and differences in their defensive mechanisms against the southern pine beetle,Dendroctonus frontalis Zimmermann, and its fungal associate,Ceratocystis minor (Hedgecock) Hunt. Both species responded to wounding and fungal inoculation by forming a hypersensitive lesion around the wound site. There were significantly less soluble sugars and more monoterpenes in the lesion tissue than in unwounded inner bark. The two species were similar in resin flow rate and inner bark soluble sugar content, but the loblolly pines had thicker bark, longer hypersensitive lesions, and a higher concentration of inner bark monoterpenes. Inner bark monoterpene composition was also significantly different between the two pine species. It is hypothesized that two different defensive strategies against southern pine beetle attack may be utilized.     

Influence of trichome exudates from species ofLycopersicon on oviposition behavior ofHeliothis zea (Boddie)

Cage experiments revealed that accessions of the wild tomato speciesLycopersicon hirsutum were preferred sites for oviposition byHeliothis zea. Hexane extracts from the leaves ofL. hirsutum were also preferred sites of oviposition in choice experiments among extracts from severalLycopersicon species. Extracts ofL. hirsutum were still biologically active several days after application, indicating that the phytochemical(s) involved are relatively stable and of low volatility. Gas Chromatographic analysis of leaf hexane extracts from 12 different accessions of theL. hirsutum complex and three tomato cultivars revealed substantial qualitative and quantitative variation in the chemical composition of these extracts. Comparison of these results with extract oviposition studies implicate a group of structurally related compounds as the active agents. Mass spectroscopy has tentatively identified these compounds as sesquiterpenes with the chemical formula C₁₅H₂₂O₂. These compounds are apparently synthesized and secreted from glandular trichomes on the leaf surface. These phytochemicals did not stimulate ovipositional behavior in females of the cabbage looper,Trichoplusia ni. The existence of genetic variation for the presence and amount of kairomones that serve as cues for insect orientation and oviposition could be utilized in a breeding program to develop tomato cultivars with genetically modified allelochemic profiles that would disrupt the sequential behavioral processes of insect host-plant selection.