Decline in gypsy moth (Lymantria dispar) performance in an elevated CO2 atmosphere depends upon host plant species

Plant species differ broadly in their responses to an elevated CO₂ atmosphere, particularly in the extent of nitrogen dilution of leaf tissue. Insect herbivores are often limited by the availability of nutrients, such as nitrogen, in their host plant tissue and may therefore respond differentially on different plant species grown in CO₂-enriched environments. We reared gyspy moth larvae (Lymantria dispar) in situ on seedlings of yellow birch (Betula allegheniensis) and gray birch (B. populifolia) grown in an ambient (350 ppm) or elevated (700 ppm) CO₂ atmosphere to test whether larval responses in the elevated CO₂ atmosphere were species-dependent. We report that female gypsy moths (Lymantria dispar) reared on gray birch (Betula populifolia) achieved similar pupal masses on plants grown at an ambient or an elevated CO₂ concentration. However, on yellow birch (B. allegheniensis), female pupal mass was 38% smaller on plants in the elevated-CO₂ atmosphere. Larval mortality was significantly higher on yellow birch than gray birch, but did not differ between the CO₂ treatments. Relative growth rate declined more in the elevated CO₂ atmosphere for larvae on yellow birch than for those on gray birch. In preference tests, larvae preferred ambient over elevated CO₂-grown leaves of yellow birch, but showed no preference between gray birch leaves from the two CO₂ atmospheres. This differential response of gypsy moths to their host species corresponded to a greater decline in leaf nutritional quality in the elevated CO₂ atmosphere in yellow birch than in gray birch. Leaf nitrogen content of yellow birch dropped from 2.68% to 1.99% while that of gray birch leaves only declined from 3.23% to 2.63%. Meanwhile, leaf condensed tannin concentration increased from 8.92% to 11.45% in yellow birch leaves while gray birch leaves only increased from 10.72% to 12.34%. Thus the declines in larval performance in a future atmosphere may be substantial and host-species-specific.     

Climate suitability and management of the gypsy moth invasion into Canada

The gypsy moth has become established throughout southern Canada east of Lake Superior where the climate is suitable for the completion of its univoltine life cycle. The spread of the gypsy moth to the north and west in Canada has so far been prevented by climatic barriers and host plant availability as well as by aggressive eradication of incipient populations. Climate change is expected to increase the area of climatic suitability and result in greater overlap with susceptible forest types throughout Canada, especially in the west. At the same time, the gypsy moth is spreading west in the USA into states bordering western Canadian provinces. These circumstances all lead to a greatly increased risk of further invasion into Canadian forests by the gypsy moth. Management actions need to be intensified in different ways in different parts of the country to reduce the impacts of spread in eastern Canada and to prevent the gypsy moth from invading western regions.     

Spatial analysis of harmonic oscillation of gypsy moth outbreak intensity

Outbreaks of many forest-defoliating insects are synchronous over broad geographic areas and occur with a period of approximately 10 years. Within the range of the gypsy moth in North America, however, there is considerable geographic heterogeneity in strength of periodicity and the frequency of outbreaks. Furthermore, gypsy moth outbreaks exhibit two significant periodicities: a dominant period of 8–10 years and a subdominant period of 4–5 years. In this study, we used a simulation model and spatially referenced time series of outbreak intensity data from the Northeastern United States to show that the bimodal periodicity in the intensity of gypsy moth outbreaks is largely a result of harmonic oscillations in gypsy moth abundance at and above a 4 km² scale of resolution. We also used geographically weighted regression models to explore the effects of gypsy moth host-tree abundance on the periodicity of gypsy moths. We found that the strength of 5-year cycles increased relative to the strength of 10-year cycles with increasing host tree abundance. We suggest that this pattern emerges because high host-tree availability enhances the growth rates of gypsy moth populations.     

Carbon and nitrogen mineralization from decomposing gypsy moth frass

Defoliation of forests by insects is often assumed to produce a pulse of available nitrogen (N) from the decomposition of frass pellets. In this study we measured rates of carbon (C) and N mineralization from gypsy moth frass incubated with and without soil, and for soil alone. Incubations were at constant temperature and soil moisture conditions and lasted for 120 days. We found that gypsy moth frass contains much labile C as well as extractable N, and that the stimulation of microbial growth by the labile C results in immobilization of essentially all of the extractable N in the frass. The response of the microbes is fast, beginning within 1 day and lasting at least 90 days. This immobilization response represents an efficient mechanism for conserving N within a forest ecosystem after a defoliation event.     

Correlation of greenhouse bioassay with field response to fertilizer by paper birch

Summary Bulk soil samples were collected from the top 15 cm of untreated areas adjacent to field fertilizer trials at 2 locations. Amounts of N, P, K, and lime equivalent to the field treatments were mixed with the soil in 15-cm diameter pots. Paper birch (Betula papyrifera March.) trees were grown from seed for a greenhouse bioassay. Height and dry weight of the bioassay seedlings were significantly correlated with 3-year volume growth of 10-year-old paper birch seedlings in the field. Correlation coefficients were 0.88 for height growth and 0.91 for dry weight growth on one site, and 0.72 and 0.63 on the other. With further refinements and observations on a larger number of sites, this bioassay technique should be a valuable tool for estimating potential response to fertilizer by young paper birch in the field, and for ranking the relative productivity of different soils.     

Hostplant,larval age,and feeding behavior influence midgut pH in the gypsy moth (Lymantria dispar)

Summary The midgut pH of late instar gypsy moth (Lymantria dispar L.) larvae is strongly alkaline, and varies with diet, larval stadium, and time since feeding. Midgut pH rises with time since feeding, and does so more quickly, reaching greater maximum values, on some diets than others. Leaf tissues of 23 tree species resist increases in alkalinity differentially; this trait and differing initial leaf pH may explain the impact of diet on gut pH. Third instar larvae may have gut conditions favorable for tannin-protein binding shortly after ingesting certain foods, but with time midgut alkalinity becomes great enough to dissociate tannin-protein complexes. Older instars rarely exhibit gut pHs low enough to permit tannin activity. Alkaline gut conditions may explain the gypsy moth's ability to feed on many tanniniferous plant species, especially in later instars. Consequences for pathogen effectiveness are discussed.     

Effects of transferring gypsy moth, Lymantria dispar, larvae between artificial diet and Quercus rubra foliage

Fourth instar gypsy moth larvae, Lymantria dispar, from a laboratory colony were transferred between artificial diet and three ages of leaves from a preferred tree, Quercus rubra. Mortality after transfers was generally minimal (<13.3%) and did not differ from mortality of larvae remaining on the same food. Larvae that were transferred to a new food accepted artificial diet and the early season foliage that is normally eaten by early instars as readily as larvae that were not transferred. Pupal weights from larvae continually fed artificial diet or transferred from artificial diet to early season foliage were not significantly different. For trials beginning during June and later, 12-day weight gain and pupal weight were greater for individuals transferred to or reared continually on artificial diet than for individuals transferred to or reared continually on mature foliage.First instars of wild larvae fed artificial diet gained less weight after 3 weeks than foliage-fed first instars. Pupal weights of wild larvae transferred from June foliage to artificial diet as fourth instars were not significantly different from pupal weights of wild larvae remaining on foliage. Thus, the response of laboratory colony larvae to artificial diet was superior to that of wild larvae.

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Résumé Des chenilles du quatrième stade de L. dispar, élevées au laboratoire sur régime artificiel et sur feuilles de différents stades de leur chêne préféré, Q. rubra, ont été transférées de l'un à l'autre de ces régimes alimentaires. La mortalité après transfert est généralement faible (<13,3%) et de même importance que pour les chenilles maintenues sur le régime artificiel. Les chenilles transférées sur un nouvel aliment ont accepté le régime artificiel ou le feuillage jeune,- normalement consommé par les premiers stades larvaires-, aussi facilement que les chenilles qui n'avaient pas été transférées. Les poids des chrysalides des chenilles élevées continuellement sur régime artificiel ou transférées sur feuillage jeune n'ont pas présenté de différences significatives. Pour des expériences commencées en juin ou ultérieurement, les gains de poids à 12 jours et pour les chrysalides sont plus élevés pour les insectes maintenus ou transférés sur régime artificiel que pour ceux transférés ou maintenus sur feuillage âgé.Le poids des chrysalides, provenant de chenilles de la nature tranférées au quatrième stade, d'un feuillage de juin à un régime artificiel, n'a pas différé significativement du poids des chrysalides provenant de chenilles maintenues sur feuillage. Les premiers stades de chenilles de la nature élevées sur régime artificiel ont pris moins de poids en trois semaines que les premiers stades ayant consommé du feuillage.

     

Effects of maternal nutrition and egg provisioning on parameters of larval hatch,survival and dispersal in the gypsy moth,Lymantria dispar L.

North American gypsy moths disperse as newly hatched larvae on wind currents in a behavior called ballooning. Because ballooning occurs before neonates begin to feed, resources used in dispersal are limited to those carried over from the egg. We show that nutritional experience of the maternal parent can influence the tendency of offspring to disperse, and that resource provisioning of eggs by the maternal parent affects the duration of the window for disperal. Offspring of females from defoliated sites had a lower tendency to balloon in a wind tunnel than larvae from females which had not experienced nutritional stress associated with host defoliation. The number of eggs in an egg mass, a reflection of the maternal parent's nutritional experience, also contributed to the predictive model for dispersal that included defoliation level. Egg weight and the levels of two yolk proteins, vitellin (Vt) and glycine-rich protein (GRP), however, had no influence of the proportion of ballooning larvae. The length of survival without food, and thus the maximum period of time for dispersal, was correlated with levels of Vt and GRP, but not with egg weight. The level of defoliation at the site from which the maternal parent was collected was not related to the longevity of offspring, nor did it have a significant effect on the levels of Vt, GRP or egg weight. Levels of hemolymph proteins arylphorin and vitellogenin in the maternal parent during the prepupal stage had no influence on levels of yolk proteins, larval longevity, or tendency to balloon.     

Simulation of litter residence times of young gypsy moth larvae and implications for predation by ants

The length of the time young gypsy moth, Lymantria dispar L. (Lepidoptera: Lymantriidae), larvae remain in litter after failing from trees would help in understanding the impact of predation by ants, as ants appear to be important predators of gypsy moths only in litter. To obtain information on caterpillar litter residence times, visual activity of larvae was investigated in the laboratory by presenting them with vertical dark silhouettes of various widths in a 1 m wide cylindrical arena. Their limit of angular resolution was about 1°. This value helped to determine distances to trees caterpillars could be expected to orient toward from randomly-located points in a forest. These data were used with results on litter crawling speed of instars 1, 2, and 3 in a simulation model that generated litter residence times averaging serveral hours. By combining results with a previously-developed model of ant predation in forest litter, it was estimated that about half of unfed first instars falling into the litter would survive, whereas only about 10% of 2nd and 3rd instars would. The implications these results have for control of the gypsy moth are discussed.

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Résumé La prédation par les fourmis du genre Formica spp. a une influence importante sur les premiers stades de L. dispar (Lepido., Lymantriidae) en Amérique du Nord. La durée du temps passé dans la litière par les jeunes chenilles après être tombées des arbres doit permettre de comprendre l'impact de la prédation par les fourmis. Pour connaître le temps de séjour des chenilles dans la litière, l'acuité visuelle des chenilles a été examinée au laboratoire en leur présentant des silhouettes noires verticales de différentes largeurs dans une arène circulaire de 1 m de diamètre. La limite de leur angle de résolution était d'environ 1°. Cette valeur a été utilisée pour déterminer les distances (jusqu'à 5 m) quelles que soient les orientations dans la forêt, d'arbres supposés susceptibles d'orienter les chenilles. Les arbres aux silhouettes acceptables étaient distribués uniformément entre 1 et 5 m.Les vitesses de déplacement dans la litière des stades 1, 2, et 3 ont été mesurées par observation directe dans la forêt (elles variaient de 0.9 à 1,54 cm/min). La distance des arbres et la vitesse de déplacement on servi à construire un modèle de simulation qui a donné un temps moyen de séjour de plusieurs heures. En combinant ces résultats avec un modèle précédemment publié de la prédation par les fourmis dans la litière de la forêt, qui utilisait le temps de contact des chenilles avec les fourmis et les taux de captures victorieuses pour estimer la mortalité, nous avons déterminé que la moitié environ des chenilles non alimentées de premier stade tombées dans la litière pourraient survivre, tandis que la survie pour le second et le troisième stade ne serait que de 10%. La discussion porte sur l'importance de ces résultats dans la lutte contre L. dispar.

     

Emigration and spatial dispersion of the gypsy moth predator Calosoma sycophanta

Mark-release studies of Calosoma sycophanta L. (Carabidae) in two 40000 m² areas were done using traps deployed in 10 by 10 grids. In both areas, gypsy moths, Lymantria dispar L. (Lymantriidae), were abundant the first but not the second year of sampling. The distribution of carabid larvae was also investigated in the first year at one of the sites and the distribution of gypsy moth larvae found under burlap bands determined for 2 years at the other site. Dispersions of beetles and prey were evaluated by Taylor's logarithmic meanvariance method and Iwao's mean-mean crowding method. Taylor's method gave the most consistent results, and the slopes of the regression lines were used to determine degree of clumping.To evaluate dispersal of adult beetles, recapture data were analysed using the Fisher-Ford and Jolly-Seber methods in conjunction with Jackson's method for separating survival and emigration. A new method for estimating emigration was found to be most consistent with the Fisher-Ford-Jackson procedure. The tendency of the beetles to aggregate (measured via Taylor's method) was generally highest for males. In one site, emigration and degree of aggregation for males appeared to vary together, while at the other site no relationship was evident. Females showed few tendencies to aggregate. Both beetle larvae and gypsy moth larvae had clumped distributions.

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Résumé L'étude par capture-recapture de Calosoma sycophanta L. (Carabidae) a été effectuée dans deux parcelles de 4 ha en utilisant des pièges distribués sur une grille de 10 sur 10. Pendant la première année d'échantillonnage, Lymantria dispar L. (Lymantriidae) était abondant et les calosomes se sont reproduits, tandis que la seconde année les papillons étaient beaucoup moins nombreux. La distribution des larves de carabes a été examinée aussi pendant la première année sur l'une des stations, et celle des chenilles pendant les deux années avec des toiles d'emballage sur l'autre station. Les dispersions des adultes et larves de calosomes et des chenilles de Lymantria ont été calculées en utilisant la régression du logarithme naturel de la variance de l'échantillon sur le logarithme naturel de la moyenne (méthode de Taylor) et la régression de l'indice de Lloyd d'agrégation moyenne sur la moyenne (méthode d'Iwao). La méthode de Taylor a donné les résultats les plus logiques, et les pentes ont servi à mesurer le degré d'agrégation. Les recaptures de calosomes adultes ont été analysées par les méthodes de Fisher-Ford et Jolly-Seber, associées à la méthode de Jackson pour séparer survie et migration. De même, une nouvelle méthode pour déterminer directement la migration en découvrant combien d'insectes recapturés ont abandonné les microparcelles s'est révélée comme la plus cohérente avec le procédé Fisher-Ford-Jackson. La tendance à l'agrégation est plus forte chez les mâles (d'après la pente du logarithme de la moyenne par rapport au logarithm e de la variance de l'échantillon). La migration et le degré d'agrégation paraissent varier ensemble dans une station, tandis que dans l'autre une telle relation n'apparaît pas clairement. Les femelles malgré des taux de migration variés mais généralement faibles, n'ont jamais pr'esenté de tendance significative à l'agrégation. A l'opposé, les larves de calosomes et de Lymantria ont presenté des distributions agrégatives.

     

Development of gypsy moth larvae feeding on red maple saplings at elevated CO<Subscript>2</Subscript> and temperature

Predicted increases in atmospheric CO₂ and global mean temperature may alter important plant-insect associations due to the direct effects of temperature on insect development and the indirect effects of elevated temperature and CO₂ enrichment on phytochemicals important for insect success. We investigated the effects of CO₂ and temperature on the interaction between gypsy moth (Lymantria dispar L.) larvae and red maple (Acer rubrum L.) saplings by bagging first instar larvae within open-top chambers at four CO₂/temperature treatments: (1) ambient temperature, ambient CO₂, (2) ambient temperature, elevated CO₂ (+300 l l^-1 CO₂), (3) elevated temperature (+3.5°C), ambient CO₂, and (4) elevated temperature, elevated CO₂. Larvae were reared to pupation and leaf samples taken biweekly to determine levels of total N, water, non-structural carbohydrates, and an estimate of defensive phenolic compounds in three age classes of foliage: (1) immature, (2) mid-mature and (3) mature. Elevated growth temperature marginally reduced (P <0.1) leaf N and significantly reduced (P <0.05) leaf water across CO₂ treatments in mature leaves, whereas leaves grown at elevated CO₂ concentration had a significant decrease in leaf N and a significant increase in the ratio of starch:N and total non-structural carbohydrates:N. Leaf N and water decreased and starch:N and total non-structural carbohydrates:N ratios increased as leaves aged. Phenolics were unaffected by CO₂ or temperature treatment. There were no interactive effects of CO₂ and temperature on any phytochemical measure. Gypsy moth larvae reached pupation earlier at the elevated temperature (female =8 days, P <0.07; male =7.5 days, P <0.03), whereas mortality and pupal fresh weight of insects were unrelated to either CO₂, temperature or their interaction. Our data show that CO₂ or temperature-induced alterations in leaf constituents had no effect on insect performance; instead, the long-term exposure to a 3.5°C increase in temperature shortened insect development but had no effect on pupal weight. It appears that in some tree-herbivorous insect systems the direct effects of an increased global mean temperature may have greater consequences for altering plant-insect interactions than the indirect effects of an increased temperature or CO₂ concentration on leaf constituents.     

Incidence and ecological relationships of pupal parasitism byBrachymeria intermedia in new Jersey populations of the gypsy moth

Pupal parasitism of the gypsy moth,Lymantria dispar (L.), was monitored in 15 study plots in New Jersey from 1978 to 1988. The predominant parasitoid was a chalcidid wasp,Brachymeria intermedia (Nees), which was found in only six plots. Parasitism was generally observed in the year of or preceding the peak numbers of gypsy moth egg masses. Parasitism exceeded 4% in only one plot. Percentage parasitism was correlated significantly with numbers of egg masses per hectare in the current season and with numbers of pupae per plot in the previous season, suggesting delayed density dependence. A multiple regression analysis found percentage parasitism to be correlated significantly with percentage infection by nuclear polyhedrosis virus, density of male host pupae, and mean minimum temperature in August and March. A canonical discriminant analysis carried out to distinguish study plots with and without the parasitoid was significant. Plots withB. intermedia had relatively higher host populations and defoliation.     

Survival and growth of eastern white pine shoot apical meristemsin vitro

Shoot apical meristems of seedling and mature eastern white pine trees were excised and grownin vitro. Placing the meristems on filters instead of directly on agarose-solidified nutrient medium enhanced survival of both juvenile and mature meristems. Applying forcing treatments to mature branches improved survival and growth of dissected meristems compared with meristems from non-forced branches in experiments conducted over two years. No consistent differences were observed among 2-, 4-, and 6-week forcing treatments. Including 5.37 nM (0.001 mg l^-1) l-naphthaleneacetic acid in the culture medium did not affect meristem survival or growth. Some meristems from seedlings grew rapidly, produced primary leaves, underwent internode elongation, and in three cases, produced adventitious roots. Meristems from mature trees did not grow as rapidly as seedling meristems. The leaves produced by mature meristems appeared to be scale leaves and a few of these had brachyblast primordia in the axils. The shoots derived from mature meristems did not produce adventitious roots.     

The effect of varying alkaloid concentrations on the feeding behavior of gypsy moth larvae, Lymantria dispar (L.) (Lepidoptera: Lymantriidae)

Nine alkaloids (acridine, aristolochic acid, atropine, berberine, caffeine, nicotine, scopolamine, sparteine, and strychnine) were evaluated as feeding deterrents for gypsy moth larvae (Lymantria dispar (L.); Lepidoptera: Lymantriidae). Our aim was to determine and compare the taste threshold concentrations, as well as the ED₅₀ values, of the nine alkaloids to determine their potency as feeding deterrents. The alkaloids were applied to disks cut from red oak leaves (Quercus rubra) (L.), a plant species highly favored by larvae of this polyphagous insect species. We used two-choice feeding bioassays to test a broad range of biologically relevant alkaloid concentrations spanning five logarithmic steps. We observed increasing feeding deterrent responses for all the alkaloids tested and found that the alkaloids tested exhibited different deterrency threshold concentrations ranging from 0.1 to 10 mM. In conclusion, it appears that this generalist insect species bears a relatively high sensitivity to these alkaloids, which confirms behavioral observations that it avoids foliage containing alkaloids. Berberine and aristolochic acid were found to have the lowest ED₅₀ values and were the most potent antifeedants. Handling Editor: Joseph Dickens.     

Influence of experience on the sequential and temporal organization of host-acceptance behavior inBrachymeria intermedia (Chalcididae), an endoparasitoid of gypsy moth

Experienced Brachymeria intermediafemales are almost twice as likely to accept a Lymantria disparpupa as inexperienced parasitoids. The sequence of parasitoid behaviors that leads to host acceptance is highly canalized. Experienced parasitoids, however, have a higher probability of initiating host investigation and making a transition from drumming to grasping, which in turn almost invariably leads to ovipositor insertion. Experienced females found the host more quickly than inexperienced females. Host-handling time did not change with experience but was longer in females that accepted rather than rejected the host. Females exposed to gypsy moth odor but not contacting pupae behaved similarly to females that never experienced host-related stimuli.     

Pheromonotropic activity in the gypsy moth Lymantria dispar: evidence for a neuropeptide

The brain-suboesophageal ganglion complex of the gypsy moth, Lymantria dispar, contains pheromonotropic activity detectable using a Helicoverpa zea in vivo bioassay for pheromone-biosynthesis-activating neuropeptide. Pheromonotropic activity was detected as early as the third larval instar and was present throughout development and through day 6 post-eclosion. Activity in the adult is presumably associated with pheromone production, while it is speculated that larval activity may be related to melanization. Adult pheromonotropic activity is associated with a peptide of approximately 3.500 kDa. It is heat labile and only partially stable when incubated at 35°C or exposed to freeze-thawing. Isolation of L. dispar pheromonotropic factor should facilitate the elucidation of the mechanism of pheromone production in this insect pest.Abbreviations ED₅₀ dose at which one-half maximal response is observal - eq equivalent - MRCH melanization and reddish colorization hormone - MW molecular weight - PBAN pheromone biosynthesis activating neuropeptide - SOG suboesophageal ganglion - TFA trifluoroacetic acid - Z11-16: Ald (Z)-11-hexadecenal     

Localization of Bacillus thuringiensis Cry1A toxin-binding molecules in gypsy moth larval gut sections using fluorescence microscopy

The microbial insecticide Bacillus thuringiensis (Bt) produces Cry toxins, proteins that bind to the brush border membranes of gut epithelial cells of insects that ingest it, disrupting the integrity of the membranes, and leading to cell lysis and insect death. In gypsy moth, Lymantria dispar, two toxin-binding molecules for the Cry1A class of Bt toxins have been identified: an aminopeptidase N (APN-1) and a 270 kDa anionic glycoconjugate (BTR-270). Studies have shown that APN-1 has a relatively weak affinity and a very narrow specificity to Cry1Ac, the only Cry1A toxin that it binds. In contrast, BTR-270 binds all toxins that are active against L. dispar larvae, and the affinities for these toxins to BTR-270 correlate positively with their respective toxicities. In this study, an immunohistochemical approach was coupled with fluorescence microscopy to localize APN-1 and BTR-270 in paraffin embedded midgut sections of L. dispar larvae. The distribution of cadherin and alkaline phosphatase in the gut tissue was also examined. A strong reaction indicative of polyanionic material was detected with alcian blue staining over the entire epithelial brush border, suggesting the presence of acidic glycoconjugates in the microvillar matrix. The Cry1A toxin-binding sites were confined to the apical surface of the gut epithelial cells with intense labeling of the apical tips of the microvilli. APN-1, BTR-270, and alkaline phosphatase were found to be present exclusively along the brush border microvilli along the entire gut epithelium. In contrast, cadherin, detected only in older gypsy moth larvae, was present both in the apical brush border and in the basement membrane anchoring the midgut epithelial cells. The topographical relationship between the Bt Cry toxin-binding molecules BTR-270 and APN-1 and the Cry1A toxin-binding sites that were confined to the apical brush border of the midgut cells is consistent with findings implicating their involvement in the mechanism of the action of Bt Cry toxins.     

Effect of dietary nitrogen on gypsy moth larval nutritional indices, development and lipid content

Nutritional indices, development rates, percent dry weights and total lipids were determined in gypsy moth larvae (Lymantria dispar L.) reared on a high wheat germ (HWG) diet or diets prepared from lyophilized, ball-milled oak or pine foliage as the only source of dietary nitrogen (N). With regard to both total and proteinaceous N content, HWG diet>oak diet>pine diet. All nutritional indices measured were significantly lower in second instars fed pine diet vs. oak diet. Protein supplementation of pine diet with either casein or ovalbumin to bring total N up to the level present in oak diet resulted in small increased in approximate digestibility (AD) and effciency of conversion of ingested food (ECI), but relative growth rate (RGR) remained unaffected. The low RGR of larvae fed pine diet (unsupplemented or protein supplemented), as compared to those fed HWG or oak diet, was accompanied by significantly lower larval percent dry weight and percent total lipid. In contrast, RGR, larval percent dry weight and total lipid values were comparable in second instars fed HWG or oak diet. Insects reared from the first through the final instar on oak diet exhibited lower pupal weights compared to those reared on HWG. Casein addition to oak diet generally resulted in even more extended larval development times and further reduced pupal weights, but wheat germ addition to oak diet did not alter development rates and caused an increase in pupal weights.     

Vertical transmission and overwintering of microsporidia in the gypsy moth, Lymantria dispar

Vertical transmission and the overwintering success of three different microsporidia infecting Lymantria dispar (Lepidoptera: Lymantriidae) larvae were investigated. Endoreticulatus schubergi, a midgut pathogen, was transmitted to offspring via female and male via the egg chorion (transovum transmission). Between 8% and 29% of the emerging larvae became infected. No spores of E. schubergi were found in surface-washed eggs. Nosema lymantriae, a microsporidium that causes systemic infections, was transovarially transmitted. Between 35% and 72% of the progeny were infected. Vairimorpha disparis, a fat body pathogen, was not vertically transmitted. The infectivity of spores that overwintered in cadavers of infected L. dispar varied by species, placement in the environment, and weather conditions. Spores of E. schubergi were still infective after an eight month exposure period of cadavers on the ground. Spores of N. lymantriae and V. disparis remained highly infective only when cadavers overwintered under a more or less continuous snow cover for four months.     

CO2 and N-fertilization effects on fine-root length, production, and mortality: a 4-year ponderosa pine study

We conducted a 4-year study of juvenile Pinus ponderosa fine root (≤2 mm) responses to atmospheric CO₂ and N-fertilization. Seedlings were grown in open-top chambers at three CO₂ levels (ambient, ambient+175 μmol/mol, ambient+350 μmol/mol) and three N-fertilization levels (0, 10, 20 g m⁻² year⁻¹). Length and width of individual roots were measured from minirhizotron video images bimonthly over 4 years starting when the seedlings were 1.5 years old. Neither CO₂ nor N-fertilization treatments affected the seasonal patterns of root production or mortality. Yearly values of fine-root length standing crop (m m⁻²), production (m m⁻² year⁻¹), and mortality (m m⁻² year⁻¹) were consistently higher in elevated CO₂ treatments throughout the study, except for mortality in the first year; however, the only statistically significant CO₂ effects were in the fine-root length standing crop (m m⁻²) in the second and third years, and production and mortality (m m⁻² year⁻¹) in the third year. Higher mortality (m m⁻² year⁻¹) in elevated CO₂ was due to greater standing crop rather than shorter life span, as fine roots lived longer in elevated CO₂. No significant N effects were noted for annual cumulative production, cumulative mortality, or mean standing crop. N availability did not significantly affect responses of fine-root standing crop, production, or mortality to elevated CO₂. Multi-year studies at all life stages of trees are important to characterize belowground responses to factors such as atmospheric CO₂ and N-fertilization. This study showed the potential for juvenile ponderosa pine to increase fine-root C pools and C fluxes through root mortality in response to elevated CO₂.