Pollen beetle offspring is more parasitized under moderate nitrogen fertilization of oilseed rape due to more attractive volatile signal.

Biocontrol providing parasitoids can orientate according to volatile organic compounds (VOCs) of their host's plants, the emission of which is potentially dependent on the availability of soil nitrogen (N). This paper aimed at finding the optimal N fertilization rate for oilseed rape (Brassica napus L.) to favor parasitism of pollen beetles (Brassicogethes aeneus Fab. syn. Meligethes aeneus Fab.) in a controlled environment. Pollen beetles preferred to oviposit into buds of plants growing under higher N fertilization, whereas their parasitoids favored moderate N fertilization. As a part of induced defense, the proportion of volatile products of glucosinolate pathway in the total oilseed rape VOC emission blend was increased. Our results suggest that the natural biological control of pollen beetle herbivory is best supported by moderate N fertilization rates.

Improved plant heat shock resistance is introduced differently by heat and insect infestation: the role of volatile emission traits.

Heat stress is one of the most important abiotic stresses confronted by plants under global climate change. Plant exposure to abiotic or biotic stress can improve its tolerance to subsequent severe episodes of the same or different stress (stress priming), but so far there is limited comparative information about how pre-exposures to different abiotic and biotic elicitors alter plant resistance to severe heat stress. We exposed the perennial herb Melilotus albus Medik., a species rich in secondary metabolites, to moderate heat stress (35 °C) and greenhouse whitefly (Trialeurodes vaporariorum West.) infestation to comparatively determine whether both pre-treatments could enhance plant tolerance to the subsequent heat shock (45 °C) stress. Plant physiological responses to stress were characterized by photosynthetic traits and volatile organic compound emissions through 72 h recovery. Heat shock treatment reduced net assimilation rate (A) and stomatal conductance in all plants, but heat-primed plants had significantly faster rates of recovery of A than other plants. By the end of the recovery period, A in none of the three heat shock-stressed groups recovered to the control level, but in whitefly-infested plants it reached the pre-heat shock level. In heat-primed plants, the heat shock treatment was associated with a fast rise of monoterpene emissions, and in whitefly-infested plants with benzenoid emissions and an increase in total phenolic content.

Volatile organic compound emissions from Alnus glutinosa under interacting drought and herbivory stresses.

Plant volatile organic compounds (VOCs) elicited in response to herbivory can serve as cues for parasitic and predatory insects, but the modification of VOC elicitation responses under interacting abiotic stresses is poorly known. We studied foliage VOC emissions in the deciduous tree Alnus glutinosa induced by feeding by the larvae of green alder sawfly (Monsoma pulveratum) under well-watered and drought-stressed conditions. Drought strongly curbed photosynthesis rate and stomatal conductance, but there were no effects of insect feeding on photosynthetic characteristics. Feeding induced emissions of volatile products of lipoxygenase pathway and monoterpenes, and emissions of stress marker compounds (E)-ß-ocimene and homoterpene DMNT. The emissions were more strongly elicited and reached a maximum value earlier in drought-stressed plants. In addition, methyl salicylate emissions were elicited in herbivory-fed drought-stressed plants. Herbivores were more strongly attracted to well-watered plants and consumed more than a four-fold greater fraction of leaf area than they consumed from drought-treated plants. Overall, this study demonstrates an important priming effect of drought, suggesting that plants under combined drought/herbivory stress are more resistant to herbivores.

Evidence for the higher importance of signal size over body size in aposematic signaling in insects.

To understand the evolution of warning coloration, it is important to distinguish between different aspects of conspicuous color patterns. As an example, both pattern element size and body size of prey have been shown to enhance the effectiveness of warning signals. However, it is unclear whether the effect of body size is merely a side effect of proportionally increasing pattern elements, or if there is an effect of body size per se. These possibilities were evaluated by offering different sized artificial caterpillars with either fixed or proportionally increasing aposematic color signal elements to wild great tits, Parus major L. (Passeriformes: Paridae). The birds' hesitation time to attack each "caterpillar" was used as a measure of the warning effect. The hesitation time showed a significant, positive size-dependence with the caterpillars whose pattern elements increased proportionally with their body size. In contrast, no size dependence was found in the larvae with fixed-size signal elements. Such a difference in mortality curves is consistent with the idea that pattern element size is a more important aspect than body size in enhancing a warning signal. Since no evidence of an effect of body size per se on signal efficiency was found, this study does not support the hypothesis that aposematic insects gain more from large size than cryptic ones.

Volatile emissions from Alnus glutionosa induced by herbivory are quantitatively related to the extent of damage.

Plant volatile organic compounds (VOCs) elicited in response to herbivory serve as cues for parasitic and predatory insects. Knowledge about quantitative relationships between the extent of herbivore-induced damage and the quantities of VOCs released is scarce. We studied the kinetics of VOC-emissions from foliage of the deciduous tree Alnus glutinosa induced by feeding activity of larvae of the geometrid moth Cabera pusaria. Quantitative relationships between the intensity of stress and strength of plant response were determined. Intensity of biotic stress was characterized by herbivore numbers (0-8 larvae) and by the amount of leaf area eaten. The strength of plant response was characterized by monitoring (i) changes in photosynthesis, (ii) leaf ultrastructure, and (iii) plant volatiles. Net assimilation rate displayed compensatory responses in herbivore-damaged leaves compared with control leaves. This compensatory response was associated with an overall increase in chloroplast size. Feeding-induced emissions of products of the lipoxygenase pathway (LOX products; (E)-2-hexenal, (Z)-3-hexenol, 1-hexanol, and (Z)-3-hexenyl acetate) peaked at day 1 after larval feeding started, followed by an increase of emissions of ubiquitous monoterpenes peaking on days 2 and 3. The emission of the monoterpene (E)-ß-ocimene and of the nerolidol-derived homoterpene 4,8-dimethyl-nona-1,3,7-triene (DMNT) peaked on day 3. Furthermore, the emission kinetics of the sesquiterpene (E,E)-α-farnesene tended to be biphasic with peaks on days 2 and 4 after start of larval feeding. Emission rates of the induced LOX products, of (E)-ß-ocimene and (E,E)-α-farnesene were positively correlated with the number of larvae feeding. In contrast, the emission of DMNT was independent of the number of feeders. These data show quantitative relationships between the strength of herbivory and the emissions of LOX products and most of the terpenoids elicited in response to feeding. Thus, herbivory-elicited LOX products and terpenoid emissions may convey both quantitative and qualitative signals to antagonists of the herbivores. In contrast, our data suggest that the feeding-induced homoterpene DMNT conveys the information "presence of herbivores" rather than information about the quantities of herbivores to predators and parasitoids.

Counterintuitive size patterns in bivoltine moths: late-season larvae grow larger despite lower food quality.

Within a season, successive generations of short-lived organisms experience different combinations of environmental parameters, such as temperature, food quality and mortality risk. Adult body size of e.g. insects is therefore expected to vary both as a consequence of proximate environmental effects as well as adaptive responses to seasonal cues. In this study, we examined intraspecific differences in body size between successive generations in 12 temperate bivoltine moths (Lepidoptera), with the ultimate goal to critically compare the role of proximate and adaptive mechanisms in determining seasonal size differences. In nearly all species, individuals developing late in the season (diapausing generation) attained a larger adult size than their conspecifics with the larval period early in the season (directly developing generation) despite the typically lower food quality in late summer. Rearing experiments conducted on one of the studied species, Selenia tetralunaria also largely exclude the possibility that the proximate effects of food quality and temperature are decisive in determining size differences between successive generations. Adaptive explanations appear likely instead: the larger body size in the diapausing generation may be adaptively associated with the lower bird predation pressure late in the season, and/or the likely advantage of large pupal size during overwintering.

Size-dependent predation risk in tree-feeding insects with different colouration strategies: a field experiment.

1. Body size is positively correlated with fecundity in various animals, but the factors that counterbalance the resulting selection pressure towards large size are difficult to establish. Positively size-dependent predation risk has been proposed as a selective factor potentially capable of balancing the fecundity advantage of large size. 2. To construct optimality models of insect body size, realistic estimates of size-dependent predation rates are necessary. Moreover, prey traits such as colouration should be considered, as they may substantially alter the relationship between body size and mortality risk. 3. To quantify mortality patterns, we conducted field experiments in which we exposed cryptic and conspicuous artificial larvae of different sizes to bird predators, and recorded the incidence of bird attacks. 4. The average daily mortality rate was estimated to vary between 4% and 10%. In both cryptic and conspicuous larvae, predation risk increased with prey size, but the increase tended to be steeper in the conspicuous group. No main effect of colour type was found. All the quantitative relationships were reasonably consistent across replicates. 5. Our results suggest that the size dependence of mortality risk in insect prey is primarily determined by the probability of being detected by a predator rather than by a size-dependent warning effect associated with conspicuous colouration. Our results therefore imply that warningly coloured insects do not necessarily benefit more than the cryptic species from large body size, as has been previously suggested.