The price of defence: toxins, visual signals and oxidative state in an aposematic butterfly

In a variety of aposematic species, the conspicuousness of an individual's warning signal and the quantity of its chemical defence are positively correlated. This apparent honest signalling in aposematism is predicted by resource competition models which assume that the production and maintenance of aposematic defences compete for access to antioxidant molecules that have dual functions as pigments directly responsible for colouration and in protecting against oxidative lipid damage. Here we study a model aposematic system - the monarch butterfly (Danaus plexippus) and make use of the variable phytochemistry of its larval host-plant, milkweeds (Asclepiadaceae), to manipulate the concentration of sequestered cardenolides. We test two fundamental assumptions of resource competition models: that (1) the possession of secondary defences is associated with costs in the form of oxidative lipid damage and reduced antioxidant defences; and (2) that oxidative damage or decreases in antioxidant defences can reduce the capacity of individuals to produce aposematic displays. Monarch caterpillars that sequestered the highest concentrations of cardenolides exhibited higher levels of oxidative lipid damage as adults. The relationship between warning signals, cardenolide concentrations and oxidative damage differed between the sexes. In male monarchs conspicuousness was explained by an interaction between oxidative damage and sequestration: as males sequester more cardenolides, those with high levels of oxidative damage become less conspicuous, while those that sequester lower levels of cardenolides equally invest in conspicuous with increasing oxidative damage. There was no significant effect of oxidative damage or concentration of sequestered cardenolides on female conspicuousness. Our results demonstrate physiological linkage between the production of coloration and protection from autotoxicity, that warning signals can be honest indicators of defensive capability, and that the relationships are different between the sexes.

Interactive effects of biotic stressors and provenance on chemical defence induction by holm oak (Quercus ilex)

Key Message The patterns of induced chemical defences in Quercus ilex leaves are specific to the biotic stress factor that causes them. Interactive effects between stressors depend on provenance. Abstract Quercus forests are suffering serious decline worldwide, closely linked to the consequences of climate change. The increase of biotic stressors threatens the survival of the holm oak (Quercus ilex), a dominant tree species in the Mediterranean Basin. A better understanding of its resistance mechanisms is urgently required to enable a better control of its decline. In this work, the ability of holm oaks from six Iberian provenances to respond to multiple biotic damage is studied through an analysis of their induced chemical defence patterns. Using 2016 seedlings established in a common garden trial (6 regions × 12 families/region × 7 seedlings/family × 4 treatments), biotic damage was induced at the root level (by infection with the widespread pathogen Phytophthora cinnamomi) and at the above-ground level (by mechanical defoliation). The levels of constitutive and induced total phenols, total tannins and condensed tannins were measured. Results showed that (1) the defensive chemical patterns present significant local and geographical variation, (2) survival to stress is more related to constitutive defences than induced ones, (3) the induced response is stressor-specific, and (4) there is an interactive effect amongst stressors whose sign (induction/inhibition) depends on the provenance. These findings on biotic stressor effects on the chemical defences and survival of holm oak can contribute to the development of genetic material selection programs in the integrated control of the widespread decline of Quercus.

Shifting chemical defence or novel weapons? A review of defence traits in Agarophyton vermiculophyllum and other invasive seaweeds

Seaweed bioinvasions increasingly affect coastal environments around the world, which increases the need for predictive models and mitigation strategies. The biotic interactions between seaweed invaders and invaded communities are often considered a key determinant of invasion success and failure and we here revise the current evidence that the capacity of seaweed invaders to deter enemies in newly reached environments correlates with their invasion success. Particularly efficient chemical defences have been described for several of the more problematic seaweed invaders during the last decades. However, confirmed cases in which seaweed invaders confronted un-adapted enemies in newly gained environments with deterrents that were absent from these environments prior to the invasion (so-called “novel weapons”) are scarce, although an increasing number of invasive and non-invasive seaweeds are screened for defence compounds. More evidence exists that seaweeds may adapt defence intensities to changing pressure by biological enemies in newly invaded habitats. However, most of this evidence of shifting defence was gathered with only one particular model seaweed, the Asia-endemic red alga Agarophyton vermiculophyllum, which is particularly accessible for direct comparisons of native and non-native populations in common garden experiments. A. vermiculophyllum interacts with consumers, epibionts and bacterial pathogens and in most of these interactions, non-native populations have rather gained than lost defensive capacity relative to native conspecifics. The increases in the few examined cases were due to an increased production of broad-spectrum deterrents and the relative scarcity of specialized deterrents perhaps reflects the circumstance that seaweed consumers and epibionts are overwhelmingly generalists.

Toxins and Other Bioactive Metabolites in Deep Chlorophyll Layers Containing the Cyanobacteria Planktothrix cf. isothrix in Two Georgian Bay Embayments, Lake Huron

The understanding of deep chlorophyll layers (DCLs) in the Great Lakes—largely reported as a mix of picoplankton and mixotrophic nanoflagellates—is predominantly based on studies of deep (>30 m), offshore locations. Here, we document and characterize nearshore DCLs from two meso-oligotrophic embayments, Twelve Mile Bay (TMB) and South Bay (SB), along eastern Georgian Bay, Lake Huron (Ontario, Canada) in 2014, 2015, and 2018. Both embayments showed the annual formation of DCLs, present as dense, thin, metalimnetic plates dominated by the large, potentially toxic, and bloom-forming cyanobacteria Planktothrix cf. isothrix. The contribution of P. cf. isothrix to the deep-living total biomass (TB) increased as thermal stratification progressed over the ice-free season, reaching 40% in TMB (0.6 mg/L at 9.5 m) and 65% in South Bay (3.5 mg/L at 7.5 m) in 2015. The euphotic zone in each embayment extended down past the mixed layer, into the nutrient-enriched hypoxic hypolimnia, consistent with other studies of similar systems with DCLs. The co-occurrence of the metal-oxidizing bacteria Leptothrix spp. and bactivorous flagellates within the metalimnetic DCLs suggests that the microbial loop plays an important role in recycling nutrients within these layers, particularly phosphate (PO4) and iron (Fe). Samples taken through the water column in both embayments showed measurable concentrations of the cyanobacterial toxins microcystins (max. 0.4 µg/L) and the other bioactive metabolites anabaenopeptins (max. ~7 µg/L) and cyanopeptolins (max. 1 ng/L), along with the corresponding genes (max. in 2018). These oligopeptides are known to act as metabolic inhibitors (e.g., in chemical defence against grazers, parasites) and allow a competitive advantage. In TMB, the 2018 peaks in these oligopeptides and genes coincided with the P. cf. isothrix DCLs, suggesting this species as the main source. Our data indicate that intersecting physicochemical gradients of light and nutrient-enriched hypoxic hypolimnia are key factors in supporting DCLs in TMB and SB. Microbial activity and allelopathy may also influence DCL community structure and function, and require further investigation, particularly related to the dominance of potentially toxigenic species such as P. cf. isothrix.

Altered capsaicin levels in domesticated chili pepper varieties affect the interaction between a generalist herbivore and its ectoparasitoid

Plant domestication has commonly reduced levels of secondary metabolites known to confer resistance against insects. Chili pepper is a special case because the fruits of different varieties have been selected for lower and higher levels of capsaicin, the main compound associated with defence. This may have important consequences for insect herbivores and their natural enemies. Despite the widespread consumption of chili peppers worldwide, the effects of capsaicin on insects are poorly understood. Here, we investigated the effect of capsaicin on a generalist herbivore, Spodoptera latifascia (Lepidoptera: Noctuidae) and its ectoparasitoid, Euplectrus platyhypenae (Hymenoptera: Eulophidae). Using chili varieties with three pungency levels: non-pungent (Padron), mild (Cayenne) and highly pungent (Habanero), as well as artificial diets spiked with three different levels of synthetic capsaicin, we determined whether higher capsaicin levels negatively affect the performance of these insects. Overall, capsaicin had a negative effect on both herbivore and parasitoid performance, particularly at high concentrations. Caterpillars reared on highly pungent fruits and high-capsaicin diet had longer development time, reduced pupation success, lower adult emergence, but also lower parasitism rates than caterpillars reared on mild or non-capsaicin treatments. In addition, we found that the caterpillars were capable of sequestering capsaicinoids in their haemolymph when fed on the high pungent variety with consequences for parasitoids’ performance and oviposition decisions. These results increase our understanding of the role of capsaicin as a chemical defence against insects and its potential implications for pest management.

Assessing the Role of Developmental and Environmental Factors in Chemical Defence Variation in Heliconiini Butterflies

Chemical defences in animals are both incredibly widespread and highly diverse. Yet despite the important role they play in mediating interactions between predators and prey, extensive differences in the amounts and types of chemical compounds can exist between individuals, even within species and populations. Here we investigate the potential role of environment and development on the chemical defences of warningly coloured butterfly species from the tribe Heliconiini, which can both synthesize and sequester cyanogenic glycosides (CGs). We reared 5 Heliconiini species in captivity, each on a single species-specific host plant as larvae, and compared them to individuals collected in the wild to ascertain whether the variation in CG content observed in the field might be the result of differences in host plant availability. Three of these species were reared as larvae on the same host plant, Passiflora riparia, to further test how species, sex, and age affected the type and amount of different defensive CGs, and how they affected the ratio of synthesized to sequestered compounds. Then, focusing on the generalist species Heliconius numata, we specifically explored variation in chemical profiles as a result of the host plant consumed by caterpillars and their brood line, using rearing experiments carried out on two naturally co-occurring host plants with differing CG profiles. Our results show significant differences in both the amount of synthesized and sequestered compounds between butterflies reared in captivity and those collected in the field. We also found a significant effect of species and an effect of sex in some, but not all, species. We show that chemical defences in H. numata continue to increase throughout their life, likely because of continued biosynthesis, and we suggest that variation in the amount of synthesized CGs in this species does not appear to stem from larval host plants, although this warrants further study. Interestingly, we detected a significant effect of brood lines, consistent with heritability influencing CG concentrations in H. numata. Altogether, our results point to multiple factors resulting in chemical defence variation in Heliconiini butterflies and highlight the overlooked effect of synthesis capabilities, which may be genetically determined to some extent.

Sex-specific differences in Juniperus communis L.: essential oil yield, growth-defence conflict and population sex-ratio

In plants, biomass and nutrient allocation often generate trade-offs between the different biochemical pathways conflicting the utilization of the common source among growth, reproduction and chemical defence. However, in dioecious plant species, these trade-off patterns could appear as a more contrasted problem between males and females due to the dissimilar reproduction investment. Generally, the growth ratio is higher in males than females, while females have a stronger defence than males. To understand the possible role of the sex-specific dissimilarities within the growth-defence conflict framework, we investigated the possible causes of the high variance of the essential oil yield in a dioecious evergreen species, Juniperus communis. Specifically, we tested the correlations between the essential oil yield with other individual-specific traits (e.g., sex, age), the presence of the growth-defence trade-off, and the differential growth and survival patterns between males and females through an extensive field-survey with sample collection in three natural populations (Kiskunság National Park, Hungary). The individual-specific essential oil yield was also measured and served as a proxy to describe the degree of chemical defence. We found that the essential oil yield showed strong and consistent sex-specific patterns decreasing with age in adults. Contrary to the predictions, the males showed a consistently higher yield than the females. We also observed a growth-defence trade-off in males but not in females. Consistently with the growth-defence conflict hypothesis, the populations’ sex-ratio was male-biased, and this pattern was more evident with ageing modifying the demographic structure due to the sexually dissimilar lifespan. Our juniper study revealed a contrasting and unique essential oil accumulation driven by the complex allocation trade-off mechanisms within individuals, which could be a flexible and adaptive defence response against the increasing biotic and abiotic environmental stresses exacerbated under global climate change.

Robotic Complexes (Facilities) of Nuclear Biological Chemical Defence Corps of the Armed Forces of the Russian Federation

The lecture is intended for training specialists in higher educational establishments according to the Federal state standard «Robotics for military and special purposes» and also for training operators of robotic complexes (facilities) for military purposes in training centers and military units. The lecture addresses two study questions: 1) the history of the development and employment of robotics technology in elimination of the consequences of the Chernobyl accident; 2) function, structure, and performance characteristics of robotic complexes (facilities) for military purposes of NBC Defence Corps. Conclusion: the material presented in the lecture will broaden the horizons and knowledge of learners on the development and use of robotics in the elimination of the consequences of accidents, as well as gain knowledge of the robotic complexes (facilities) currently being supplied to the NBC Defence Corps.