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Eastern hemlock (Tsuga canadensis (L.) Carriere) is an eastern North American conifer threatened by the invasive hemlock woolly adelgid (Adelges tsugae Annand). Changes in foliar terpenes and phenolics were evaluated in new (current-year growth) and mature (1-year-old growth) hemlock needles during the growing season and into plant dormancy. From April through September, foliar concentrations of nonvolatile soluble phenolics, condensed tannins, lignin, mono- and sesquiterpenes, [alpha]-pinene, camphene, isobornyl acetate, and diterpene resin were quantified. After September, additional analyses of metabolites that continued to differ significantly between new and mature foliage were carried out. Total soluble phenolic and condensed tannin concentrations in new foliage remained low relative to those of mature foliage throughout the growing season and converged in December. Lignin concentration in new foliage converged with that of mature foliage by July. Concentrations of [alpha]-pinene, camphene, isobornyl acetate, and diterpene resin in new foliage converged with those of mature foliage within 1 month of budbreak. The convergence of terpene concentrations in new and mature foliage suggests that these metabolites may play a role in herbivore defense during the peak growing season. Conversely, soluble phenolics, including condensed tannins, may defend foliage from herbivory outside of the spring growth period. Key words: phenology, terpenes, phenolics, conifers, hemlock. La pruche du Canada (Tsuga canadensis (L.) Carriere) est un conifere de l'est de l'Amerique du Nord menace par un insecte invasif, le puceron lanigere de la pruche (Adelges tsugae Annand). Les modifications dans les composes phenoliques et les terpenes foliaires ont ete evaluees dans les nouvelles aiguilles (pousses de l'annee en cours) et les aiguilles matures (pousses agees d'un an) de la pruche durant la saison de croissance et la periode de dormance. La concentration foliaire des composes phenoliques solubles non volatiles, des tannins condenses, des mono- et sesquiterpenes, [alpha]-pinene, camphene, acetate d'isobornyle, et des diterpenes resiniques a ete quantifiee du mois d'avril au mois de septembre. Apres le mois de septembre, d'autres analyses des metabolites qui continuaient a differer significativement dans le nouveau feuillage comparativement au feuillage mature ont ete effectuees. La concentration totale de composes phenoliques solubles et de tannins condenses dans le nouveau feuillage demeurait faible comparativement au feuillage mature durant toute la saison de croissance et convergeait en decembre. La concentration de lignine dans le nouveau feuillage convergeait vers celle du feuillage mature vers le mois de juillet. La concentration de [alpha]-pinene, de camphene, d'acetate d'isobornyle et de diterpenes resiniques dans le nouveau feuillage convergeait vers celle du feuillage mature a moins d'un mois du debourrement. La convergence de la concentration de terpenes dans le nouveau feuillage et le feuillage mature indique que ces metabolites jouent possiblement un role dans la defense contre les herbivores au plus fort de la saison de croissance. A l'inverse, les composes phenoliques solubles, incluant les tannins condenses, pourraient proteger le feuillage contre les herbivores en dehors de la periode de croissance printaniere. [Traduit par la Redaction] Mots-cles: phenologie, terpenes, composes phenoliques, coniferes, pruche., Introduction Phenology is the study of seasonally varying developmental events driven by environmental cues. Plant phenology is characterized by temporal patterns of growth associated with abiotic factors such as degree-day, [...]

Lymantria dispar L. is an invasive, non-native defoliating Lepidopteran established in North America that feeds on forest and urban trees. While many products are available to manage L. dispar post-emergence, few exist to prevent egg hatch when applied to egg masses. Here, we present the results of 3 separate experiments aimed at determining the efficacy of pre-emergent insecticides against L. dispar egg hatch. We found that the labeled rate (1:1) of Golden Pest Spray Oil (GPSO; AI: 93% soybean oil) can prevent L. dispar larvae from emerging in both field and lab assays. In large public spaces, we found that this treatment was ineffective at preventing L. dispar emergence or defoliation. Acelepryn (AI: 18.4% chlorantraniliprole) resulted in some suppression of egg hatch at a very low rate (.06 ml/ 3.8 liter) in both lab and field settings and the efficacy of higher rates should be further investigated. We also tested GPSO against Lepitect (97.4% acephate) in a public area that also received a Foray 48B (12.65% Bacillus thuringiensis, subsp. kurstaki) aerial application. On large oak trees in public areas, GPSO and Lepitect were not effective at reducing defoliation. Dormant pesticide applications generally reduce the risk of affecting negatively predator and parasitoid communities and are therefore desirable. Lymantria dispar pre-egg hatch applications will not work in every situation but should be considered as part of an integrated pest management (IPM) strategy for individual homeowner trees where thorough coverage can be obtained. [ABSTRACT FROM AUTHOR]

Our understanding of how conifers respond biochemically to multiple simultaneous herbivore attacks is lacking. Eastern hemlock (Tsuga canadensis ; 'hemlock') is fed on by hemlock woolly adelgid (Adelges tsugae ; 'adelgid') and by later-instar gypsy moth (Lymantria dispar ; 'gypsy moth') caterpillars. The adelgid is a stylet-feeding insect that causes a salicylic acid (SA)-linked response in hemlock, and gypsy moth larvae are folivores that presumably cause a jasmonic acid (JA)-linked response. This system presents an opportunity to study how invasive herbivore–herbivore interactions mediated through host biochemical responses. We used a factorial field experiment to challenge chronically adelgid-infested hemlocks with gypsy moth caterpillars. We quantified 17 phytohormones, 26 phenolic and terpene metabolites, and proanthocyanidin, cell wall-bound (CW-bound) phenolic, and lignin contents. Foliage infested with adelgid only accumulated gibberellins and SA; foliage challenged by gypsy moth only accumulated JA phytohormones. Gypsy moth folivory on adelgid-infested foliage reduced the accumulation of JA phytohormones and increased the SA levels. Both herbivores increased CW-bound phenolics and gypsy moth increased lignin content when feeding alone but not when feeding on adelgid-infested foliage. Our study illustrates the importance of understanding the biochemical mechanisms and signaling antagonism underlying tree responses to multiple stresses and of disentangling local and systemic stress signaling in trees. [ABSTRACT FROM AUTHOR]

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1. Interactions between invertebrate herbivores with different feeding modes are common on long‐lived woody plants. In cases where one herbivore facilitates the success of another, the consequences for their shared host plant may be severe. Eastern hemlock (Tsuga canadensis), a canopy‐dominant conifer native to the eastern U.S., is currently threatened with extirpation by the invasive stylet‐feeding hemlock woolly adelgid (Adelges tsugae). The effect of adelgid on invasive hemlock‐feeding folivores remains unknown. 2. This study evaluated the impact of feeding by hemlock woolly adelgid on gypsy moth (Lymantria dispar) larval preference for, and performance on, eastern hemlock. To assess preference, 245 field‐grown hemlocks were surveyed for gypsy moth herbivory damage and laboratory paired‐choice bioassays were conducted. To assess performance, gypsy moth larvae were reared to pupation on adelgid‐infested or uninfested hemlock foliage, and pupal weight, proportional weight gain, and larval period were analysed. 3. Adelgid‐infested hemlocks experienced more gypsy moth herbivory than did uninfested control trees, and laboratory tests confirmed that gypsy moth larvae preferentially feed on adelgid‐infested hemlock foliage. Gypsy moth larvae reared to pupation on adelgid‐infested foliage gained more weight than larvae reared on uninfested control foliage. 4. These results suggest that the synergistic effect of adelgid and gypsy moth poses an additional threat to eastern hemlock that may increase extirpation risk and ecological impact throughout most of its range. [ABSTRACT FROM AUTHOR]

Trees attacked by multiple herbivores need to defend themselves against dynamic biotic challenges; appropriate responses to one stressor can elicit hormonal responses that are antagonistic to another. Hemlock (Tsuga canadensis) infestation by hemlock woolly adelgid (HWA; Adelges tsugae) results in the accumulation of the defensive hormone salicylic acid.We explored the potential for HWA infestation to interfere with anti‐folivore‐induced defence signalling and its implications for a native folivore (hemlock looper; Lambdina fiscellaria). Hemlocks were infested with HWA and/or sprayed with methyl jasmonate; foliar defences were analyzed and foliage quality for looper larvae was assessed.Both treatments activated foliar defensive traits, including a HWA‐mediated increase in peroxidase activity and an accumulation of cell wall‐bound phenolics and lignin, as well as a methyl jasmonate‐mediated increase in lipoxygenase activity. The two treatments had an additive effect on other defensive traits and both treatments negatively affected looper performance.These results suggest that salicylic acid and jasmonic acid are not strictly antagonistic in conifers and that both have a role in anti‐folivore defence signalling. The present study illustrates the need for a better understanding of hormone signalling, cross‐talk and induced responses in conifers. [ABSTRACT FROM AUTHOR]

In its native range in Asia, emerald ash borer (EAB; Agrilus planipennis Fairmaire) is a secondary colonizer of its coevolved hosts, including Manchurian ash (Fraxinus mandschurica Ruprecht). However, why larval performance is enhanced on stressed trees remains to be determined.We examined the mechanisms that may reduce the resistance of stressed Manchurian ashes. Trees in a common garden were girdled or left untreated as controls and both treatments were inoculated with EAB eggs. Larvae and bark tissue were harvested after feeding and larval performance so that the defensive mechanisms and nutritional quality of bark tissue could be assessed.Larval survival and mass was twice as high on girdled trees, although girdling had no effect on bark phenolics. The activity of two enzymes involved in wound repair and lignification (peroxidase and cinnamic alcohol dehydrogenase) was increased by girdling in bark tissue. Starch and total protein levels declined in girdled stems, although sugar content was unchanged. Total disulphide levels, a measure of protein oxidative damage, were increased by girdling, although no difference in lipid oxidative damage in ash phloem tissue was detected.Girdling ash increased larval performance even though it elevated some wound repair mechanisms and decreased some indices of plant nutritional quality. Other tree responses to girdling also were poor predictors of larval performance. It appears that mechanisms of stress‐induced variation in resistance of Manchurian ash to EAB differ from those that may explain interspecific variation in resistance. [ABSTRACT FROM AUTHOR]

Predators that feed on chemically-defended prey often experience non-lethal effects that result in learned avoidance of the prey species. Some predators are able to consume toxic prey without ill-effect. The Chinese mantid is able to consume cardenolide-containing monarch caterpillars without immediate adverse effects. Although they discard the caterpillars' gut contents, mantids consume sequestered cardenolides. Although consumption of these cardenolides does not elicit an acute response, there may be long-term costs associated with cardenolide consumption. We tested the hypothesis that consumption of monarch caterpillars will adversely affect adult mantid biomass gain and reproductive condition. We reared mantids from egg to adult and assigned them to one of four toxicity groups that differed in the number of monarch caterpillars offered over a 15-day period. Mantids consumed similar amounts of prey biomass during the experiment. Yet, mantids in the high-toxicity group had a higher conversion efficiency and gained more biomass than mantids in other groups. Mantids in all treatment groups produced similar numbers of eggs. However, mantids in the high-toxicity group produced heavier eggs and devoted a greater portion of their biomass toward egg production than those in the control group. This increase in reproductive condition is probably driven by variation in prey nutritional value and/or the nutritional advantages inherent in eating multiple food types. Our results demonstrate the mantid is able to incorporate 'toxic' monarch prey into its diet without acute or chronic ill-effects. [ABSTRACT FROM AUTHOR]

The "mother knows best" hypothesis states that adults should choose hosts for oviposition on which their offspring will best perform, maximizing their own fitness. It has been hypothesized that this preference-performance relationship for wood-boring insects is especially important because larvae are not able to switch hosts, although no study has examined oviposition choices for these insects. We examined oviposition preferences of the emerald ash borer, Agrito planipennis Fairmaire (Coleoptera: Buprestidae), in two common gardens, one on the campus of Wright State University in Dayton, OH, and the other at the Michigan State University Tollgate Research Farm in Novi, MI, by wrapping cheesecloth around ash trunks to assess passive oviposition patterns. We found that in both gardens, ash species native to North America, which are highly susceptible to the emerald ash borer, consistently received more ova than Manchurian ash, which is indigenous to Asia and more resistant to the emerald ash borer. Susceptible trees in the Novi garden received 93 times the number of ova and susceptible trees at the Wright State garden received up to 25 times the number of ova that were received by Manchurian ash in each of their respective gardens. Neither tree size nor vigor affected oviposition choice. There were also higher numbers of adult exit holes on North American than Manchurian ash in both common gardens. The observed oviposition preferences in this study align with patterns of adult feeding preference, ash host mortality, and exit hole numbers from other studies. These observations also suggest that oviposition preferences may contribute to interspecific patterns of host resistance and mortality. Collectively, our results demonstrate that the emerald ash borer prefers to oviposit on species on which its offspring will best perform, suggesting that there is strong selection for the ability to recognize host cues that predict better larval survival and performance. [ABSTRACT FROM AUTHOR]

The emerald ash borer, Agrilus planipennis Fairmaire, has destroyed millions of ash trees (Fraxinus spp.) in North America since first identified in Detroit in 2002. With species of ash distributed throughout North America, it is easy to speculate the extinction of all susceptible species of ash on the continent given a lack of physical, environmental, or climactic barrier for dispersal of the insect. We investigated water balance characteristics of emerald ash borer ova by using gravimetric methods in an effort to measure their response to heat- and water-stress and explore possible influences this stress may have on the ecology and physiology of the ovum. We also explored the possible water balance benefit of a peculiar, "clustering," oviposition behavior, as well as the difference in responses to stress between ova from a laboratory colony and ova from two wild populations. We found no evidence of water vapor absorption as a water balance strategy; rather enhanced water retention, resistance to desiccation, and viability with low water content were important survival strategies for these ova. Surface lipids resist thermal breakdown as indicated by ova having no detectable critical transition temperature, maintaining their water-proofing function as temperature rises. The observed "clustering" behavior had no desiccation-avoidance benefit and ova from the wild populations behaved almost identically to the ova from the lab colony, although the lab ova were slightly larger and more sensitive to dehydration. Given this new information, there appears to be no heat- or water-stress barriers for the dispersal of this devastating pest at the ovum stage. [ABSTRACT FROM AUTHOR]

1. Chronic nitrogen (N) deposition may alter the bioavailability of dissolved organic matter (DOM) in streams by multiple pathways. Elevated N deposition may alter the nutrient stoichiometry of DOM as well as nutrient availability in stream water. 2. We evaluated the influence of a decadal-scale experimental N enrichment on the relative importance of DOM nutrient content and inorganic nutrient availability on the bioavailability of DOM. We measured the consumption of dissolved organic carbon (DOC) and changes in nutrient concentration, DOM components and enzyme activity in a bottle incubation assay with different DOM and nutrient treatments. To evaluate the effect of DOM stoichiometry, we used leaf leachates of different carbon/N/phosphorus (C : N :P) ratio, made from leaf litter sourced in the reference and N-enriched catchments at the Bear Brook Watershed in Maine (BBWM). We also manipulated the concentration of inorganic N and P to compare the effect of nutrient enrichment with DOM stoichiometry. 3. DOC from the N-enriched catchment was consumed 14% faster than that from the reference catchment. However, mean DOC consumption for both leachates was more than doubled by the simultaneous addition of N and P, compared to controls, while the addition of N or P alone increased consumption by 42 and 23%, respectively. The effect of N and/or P enrichment consistently had a greater effect than DOM source for all response variables considered. 4. We subsequently conducted DOC uptake measurements using leaf leachate addition under ambient and elevated N and P in the streams draining the reference and N-enriched catchments at BBWM. In both streams, DOC uptake lengths were shorter when N and P were elevated. 5. Although both DOM stoichiometry and inorganic nutrient availability affect DOM bioavailability, N and P co-limitation appears to be the dominant driver of reach-scale processing of DOM. [ABSTRACT FROM AUTHOR]

Features of Balaustium sp. include resistance to intense heat and desiccation, affinity for hot surfaces in bright light, abundance in semi-arid/arid biotopes, and a large pair of secretory glands called umulae with no known function (defense secretion excepted). Here we show that the urnulae secrete a waterproofing barrier that reduces the mite's cuticular permeability to water. Exposure to white light was used to stimulate release of the secretion; the urnulae protruded and exuded streams of red fluid at the tip of this structure that covered the entire body. Results showed that mites coated with urnulae secretion lost water at approximately half the rate of mites that did not secrete. Similarly, urnulae secretion coated mites demonstrated an increase in water-tightness of the cuticle reflected by a 9°C elevation in temperature threshold for water loss on an evaporation curve, increasing their optimal temperature tolerance for survival (lethal permeability temperature, LPT). Results also show a 10 kJ/mol drop in activation energy (Ea) for water loss, representative of a substantial cuticular modification, and a decrease in Arrhenius frequency steric factor A, indicating an overall decrease in body water losses. The absence of a critical transition temperature (CTT), however, reveals that urnulae secretion coating functions to resist a phase change as the temperature rises, permitting the mites to cope with high temperature without succumbing to water and heat stress, by inhibiting cuticular breakdown. [ABSTRACT FROM AUTHOR]