Many parasitoids lack adult fat accumulation, despite fatty acid synthesis: A discussion of concepts and considerations for future research.

Fat reserves, specifically the accumulation of triacylglycerols, are a major energy source and play a key role for life histories. Fat accumulation is a conserved metabolic pattern across most insects, yet in most parasitoid species adults do not gain fat mass, even when nutrients are readily available and provided ad libitum. This extraordinary physiological phenotype has evolved repeatedly in phylogenetically dispersed parasitoid species. This poses a conundrum because it could lead to significant constraints on energy allocation toward key adult functions such as survival and reproduction. Recent work on the underlying genetic and biochemical mechanisms has spurred a debate on fat accumulation versus fat production, because of incongruent interpretation of results obtained using different methodologies. This debate is in part due to semantics, highlighting the need for a synthetic perspective on fat accumulation that reconciles previous debates and provides new insights and terminology. In this paper, we propose updated, unambiguous terminology for future research in the field, including "fatty acid synthesis" and "lack of adult fat accumulation", and describe the distinct metabolic pathways involved in the complex process of lipogenesis. We then discuss the benefits and drawbacks of the main methods available to measure fatty acid synthesis and adult fat accumulation. Most importantly, gravimetric/colorimetric and isotope tracking methods give complementary information, provided that they are applied with appropriate controls and interpreted correctly. We also compiled a comprehensive list of fat accumulation studies performed during the last 25 years. We present avenues for future research that combine chemistry, ecology, and evolution into an integrative approach, which we think is needed to understand the dynamics of fat accumulation in parasitoids.

Climate Change, Extreme Temperatures and Sex-Related Responses in Spiders.

Climatic extremes, such as heat waves, are increasing in frequency, intensity and duration under anthropogenic climate change. These extreme events pose a great threat to many organisms, and especially ectotherms, which are susceptible to high temperatures. In nature, many ectotherms, such as insects, may seek cooler microclimates and 'ride out´ extreme temperatures, especially when these are transient and unpredictable. However, some ectotherms, such as web-building spiders, may be more prone to heat-related mortality than more motile organisms. Adult females in many spider families are sedentary and build webs in micro-habitats where they spend their entire lives. Under extreme heat, they may be limited in their ability to move vertically or horizontally to find cooler microhabitats. Males, on the other hand, are often nomadic, have broader spatial distributions, and thus might be better able to escape exposure to heat. However, life-history traits in spiders such as the relative body size of males and females and spatial ecology also vary across different taxonomic groups based on their phylogeny. This may make different species or families more or less susceptible to heat waves and exposure to very high temperatures. Selection to extreme temperatures may drive adaptive responses in female physiology, morphology or web site selection in species that build small or exposed webs. Male spiders may be better able to avoid heat-related stress than females by seeking refuge under objects such as bark or rocks with cooler microclimates. Here, we discuss these aspects in detail and propose research focusing on male and female spider behavior and reproduction across different taxa exposed to temperature extremes.

Soil legacy effects of plants and drought on aboveground insects in native and range-expanding plant communities.

Soils contain biotic and abiotic legacies of previous conditions that may influence plant community biomass and associated aboveground biodiversity. However, little is known about the relative strengths and interactions of the various belowground legacies on aboveground plant-insect interactions. We used an outdoor mesocosm experiment to investigate the belowground legacy effects of range-expanding versus native plants, extreme drought and their interactions on plants, aphids and pollinators. We show that plant biomass was influenced more strongly by the previous plant community than by the previous summer drought. Plant communities consisted of four congeneric pairs of natives and range expanders, and their responses were not unanimous. Legacy effects affected the abundance of aphids more strongly than pollinators. We conclude that legacies can be contained as soil 'memories' that influence aboveground plant community interactions in the next growing season. These soil-borne 'memories' can be altered by climate warming-induced plant range shifts and extreme drought.

The Biology and Ecology of Parasitoid Wasps of Predatory Arthropods.

Parasitoid wasps are important components of insect food chains and have played a central role in biological control programs for over a century. Although the vast majority of parasitoids exploit insect herbivores as hosts, others parasitize predatory insects and arthropods, such as ladybird beetles, hoverflies, lacewings, ground beetles, and spiders, or are hyperparasitoids. Much of the research on the biology and ecology of parasitoids of predators has focused on ladybird beetles, whose parasitoids may interfere with the control of insect pests like aphids by reducing ladybird abundance. Alternatively, parasitoids of the invasive ladybird Harmonia axyridis may reduce its harmful impact on native ladybird populations. Different life stages of predatory insects and spiders are susceptible to parasitism to different degrees. Many parasitoids of predators exhibit intricate physiological interrelationships with their hosts, adaptively manipulating host behavior, biology, and ecology in ways that increase parasitoid survival and fitness.

Reviewing the relationship between neoliberal societies and nature: implications of the industrialized dominant social paradigm for a sustainable future.

How a society relates to nature is shaped by the dominant social paradigm (DSP): a society's collective view on social, economic, political, and environmental issues. The characteristics of the DSP have important consequences for natural systems and their conservation. Based on a synthesis of academic literature, we provide a new gradient of 12 types of human-nature relationships synthesized from scientific literature, and an analysis of where the DSP of industrialized, and more specifically, neoliberal societies fit on that gradient. We aim to answer how the industrialized DSP relates to nature, i.e., what types of human-nature relationships this DSP incorporates, and what the consequences of these relationships are for nature conservation and a sustainable future. The gradient of human-nature relationships is based on three defining characteristics: (1) a nature-culture divide, (2) core values, and (3) being anthropocentric or ecocentric. We argue that the industrialized DSP includes elements of the anthropocentric relationships of mastery, utilization, detachment, and stewardship. It therefore regards nature and culture as separate, is mainly driven by instrumental values, and drives detachment from and commodification of nature. Consequently, most green initiatives and policies driven by an industrialized and neoliberal DSP are based on economic incentives and economic growth, without recognition of the needs and limits of natural systems. This leads to environmental degradation and social inequality, obstructing the path to a truly sustainable society. To reach a more ecocentric DSP, systemic changes, in addition to individual changes, in the political and economic structures of the industrialized DSP are needed, along with a change in values and approach toward nature, long-term sustainability, and conservation.

Effects of Light Quality on Colonization of Tomato Roots by AMF and Implications for Growth and Defense.

Beneficial soil microbes can enhance plant growth and defense, but the extent to which this occurs depends on the availability of resources, such as water and nutrients. However, relatively little is known about the role of light quality, which is altered during shading, resulting a low red: far-red ratio (R:FR) of light. We examined how low R:FR light influences arbuscular mycorrhizal fungus (AMF)-mediated changes in plant growth and defense using Solanum lycopersicum (tomato) and the insect herbivore Chrysodeixis chalcites. We also examined effects on third trophic level interactions with the parasitoid Cotesia marginiventris. Under low R:FR light, non-mycorrhizal plants activated the shade avoidance syndrome (SAS), resulting in enhanced biomass production. However, mycorrhizal inoculation decreased stem elongation in shaded plants, thus counteracting the plant's SAS response to shading. Unexpectedly, activation of SAS under low R:FR light did not increase plant susceptibility to the herbivore in either non-mycorrhizal or mycorrhizal plants. AMF did not significantly affect survival or growth of caterpillars and parasitoids but suppressed herbivore-induced expression of jasmonic acid-signaled defenses genes under low R:FR light. These results highlight the context-dependency of AMF effects on plant growth and defense and the potentially adverse effects of AMF under shading.

Prey availability affects developmental trade-offs and sexual-size dimorphism in the false widow spider, Steatoda grossa.

In many spiders, females are significantly larger than males. Several theories have been postulated to explain sexual size dimorphism (SSD), including differential predation risks experienced by each sex early in life (including female cannibalism of males), male-male competition, and the more costly production of eggs than sperm. However, there is considerable intraspecific variation in the relative size of males and females that is reflected in trade-offs on traits such as growth rate and body size. When SSD favors female size, the body mass ratios between the smallest and largest males is expected to be much greater than in females. Here, growth trajectories and body masses of the false widow spider, Steatoda grossa, were compared in male and female spiders fed continually or intermittently. Males provided with unlimited prey (fruit flies and house crickets) took about 15 weeks to attain full size and sexual maturity and grew to a mean of 25 mg. By contrast, males fed only once every three weeks took approximately 6 weeks longer to reach maturity but were only about half as large (mean 13 mg) as males fed constantly. Females fed intermittently took almost twice as long (45 weeks versus 24 weeks) as constantly-fed females to reach maturity, but were almost 90% as large when fully grown. These results reveal that, although both sexes trade-off development time and body size to achieve the optimal phenotype, rapid development is more important than larger body size in males whereas the opposite is true in females. This finding supports life-history theory underpinning sexual-size dimorphism in some spider lineages.

Extrinsic Inter- and Intraspecific Competition in Parasitoid Wasps.

The diverse ecology of parasitoids is shaped by extrinsic competition, i.e., exploitative or interference competition among adult females and males for hosts and mates. Adult females use an array of morphological, chemical, and behavioral mechanisms to engage in competition that may be either intra- or interspecific. Weaker competitors are often excluded or, if they persist, use alternate host habitats, host developmental stages, or host species. Competition among adult males for mates is almost exclusively intraspecific and involves visual displays, chemical signals, and even physical combat. Extrinsic competition influences community structure through its role in competitive displacement and apparent competition. Finally, anthropogenic changes such as habitat loss and fragmentation, invasive species, pollutants, and climate change result in phenological mismatches and range expansions within host-parasitoid communities with consequent changes to the strength of competitive interactions. Such changes have important ramifications not only for the success of managed agroecosystems, but also for natural ecosystem functioning.

Climate change-mediated temperature extremes and insects: From outbreaks to breakdowns.

Insects are among the most diverse and widespread animals across the biosphere and are well-known for their contributions to ecosystem functioning and services. Recent increases in the frequency and magnitude of climatic extremes (CE), in particular temperature extremes (TE) owing to anthropogenic climate change, are exposing insect populations and communities to unprecedented stresses. However, a major problem in understanding insect responses to TE is that they are still highly unpredictable both spatially and temporally, which reduces frequency- or direction-dependent selective responses by insects. Moreover, how species interactions and community structure may change in response to stresses imposed by TE is still poorly understood. Here we provide an overview of how terrestrial insects respond to TE by integrating their organismal physiology, multitrophic, and community-level interactions, and building that up to explore scenarios for population explosions and crashes that have ecosystem-level consequences. We argue that TE can push insect herbivores and their natural enemies to and even beyond their adaptive limits, which may differ among species intimately involved in trophic interactions, leading to phenological disruptions and the structural reorganization of food webs. TE may ultimately lead to outbreak-breakdown cycles in insect communities with detrimental consequences for ecosystem functioning and resilience. Lastly, we suggest new research lines that will help achieve a better understanding of insect and community responses to a wide range of CE.

Detoxification of plant defensive glucosinolates by an herbivorous caterpillar is beneficial to its endoparasitic wasp.

Plant chemical defences impact not only herbivores, but also organisms in higher trophic levels that prey on or parasitize herbivores. While herbivorous insects can often detoxify plant chemicals ingested from suitable host plants, how such detoxification affects endoparasitoids that use these herbivores as hosts is largely unknown. Here, we used transformed plants to experimentally manipulate the major detoxification reaction used by Plutella xylostella (diamondback moth) to deactivate the glucosinolate defences of its Brassicaceae host plants. We then assessed the developmental, metabolic, immune, and reproductive consequences of this genetic manipulation on the herbivore as well as its hymenopteran endoparasitoid Diadegma semiclausum. Inhibition of P. xylostella glucosinolate metabolism by plant-mediated RNA interference increased the accumulation of the principal glucosinolate activation products, the toxic isothiocyanates, in the herbivore, with negative effects on its growth. Although the endoparasitoid manipulated the excretion of toxins by its insect host to its own advantage, the inhibition of herbivore glucosinolate detoxification slowed endoparasitoid development, impaired its reproduction, and suppressed the expression of genes of a parasitoid-symbiotic polydnavirus that aids parasitism. Therefore, the detoxification of plant glucosinolates by an herbivore lowers its toxicity as a host and benefits the parasitoid D. semiclausum at multiple levels.

Range-Expansion in Processionary Moths and Biological Control.

Global climate change is resulting in a wide range of biotic responses, including changes in diel activity and seasonal phenology patterns, range shifts polewards in each hemisphere and/or to higher elevations, and altered intensity and frequency of interactions between species in ecosystems. Oak (Thaumetopoea processionea) and pine (T. pityocampa) processionary moths (hereafter OPM and PPM, respectively) are thermophilic species that are native to central and southern Europe. The larvae of both species are gregarious and produce large silken 'nests' that they use to congregate when not feeding. During outbreaks, processionary caterpillars are capable of stripping foliage from their food plants (oak and pine trees), generating considerable economic damage. Moreover, the third to last instar caterpillars of both species produce copious hairs as a means of defence against natural enemies, including both vertebrate and invertebrate predators, and parasitoids. These hairs contain the toxin thaumetopoein that causes strong allergic reactions when it comes into contact with human skin or other membranes. In response to a warming climate, PPM is expanding its range northwards, while OPM outbreaks are increasing in frequency and intensity, particularly in northern Germany, the Netherlands, and southern U.K., where it was either absent or rare previously. Here, we discuss how warming and escape from co-evolved natural enemies has benefitted both species, and suggest possible strategies for biological control.

Population- and Species-Based Variation of Webworm-Parasitoid Interactions in Hogweeds (Heracelum spp.) in the Netherlands.

In three Dutch populations of the native small hogweed (Heracleum sphondylium L. [Apiales: Apiaceae]), and one of the invasive giant hogweed (H. mantegazzianum Sommeier & Levier [Apiales: Apiaceae]), interactions between a specialist herbivore, the parsnip webworm (Depressaria radiella), and its associated parasitoids were compared during a single growing season. We found host plant species-related differences in the abundance of moth pupae, the specialist polyembryonic endoparasitoid, Copidosoma sosares, the specialist pupal parasitoid, Barichneumon heracliana, and a potential hyperparasitoid of C. sosares, Tyndaricus scaurus Walker (Hymenoptera: Encyrtidae). Adult D. radiella body mass was similar across the three small hogweed populations, but moths and their pupal parasitoid B. heracliana were smaller when developing on giant than on small hogweeds where the two plants grew in the same locality (Heteren). Mixed-sex and all-male broods of C. sosares were generally bigger than all-female broods. Furthermore, adult female C. sosares were larger than males and adult female mass differed among the three small hogweed populations. The frequency of pupal parasitism and hyperparasitism also varied in the different H. sphondylium populations. These results show that short-term (intra-seasonal) effects of plant population on multitrophic insects are variable among different species in a tightly linked food chain.

Effects of elevated CO2 and temperature on survival and wing dimorphism of two species of rice planthoppers (Hemiptera: Delphacidae) under interaction.

BACKGROUND: Anthropogenic climate change (ACC) may have significant impacts on insect herbivore communities including pests. Two of the most important climate-change related factors are increased atmospheric concentrations of carbon dioxide (CO2 ), and increasing mean global temperature. Although increasing attention is being paid to the biological and ecological effects of ACC, important processes such as interspecific interaction between insect herbivores have been little explored. Here, in a field experiment using the FACE (free-air CO2 enrichment) system, we investigated the effect of elevated CO2 and temperature on survival and wing dimorphism of two species of rice planthoppers, Laodelphax striatellus and Nilaparvata lugens under interaction. RESULTS: The two species were grouped into five treatments of relative density (0/50, 13/37, 25/25, and 37/13, 50/0), each of which was allocated to one of a factorial combination of two CO2 concentrations and two temperature treatments (elevated and ambient levels). Our results revealed that climatic treatment has no effects on survivorship of interspecific competing planthoppers. However, climatic treatment affected wing-form of planthoppers under interspecific interaction. For females of N. lugens, in the 37/13 ratio, proportion macropterours form was lower under elevated CO2 + temperature than under the ambient environment or than under elevated temperature. For females of L. striatellus, proportion macropterous form did not differ among climatic treatments at each ratio treatment. CONCLUSION: These findings illustrate that climate change-related factors, by affecting the macropetry of interspecific competing planthoppers, may influence planthopper fitness. We provide new information that could assist with forecasting outbreaks of these migratory pests. © 2020 Society of Chemical Industry.

The ecological role of bacterial seed endophytes associated with wild cabbage in the United Kingdom.

Endophytic bacteria are known for their ability in promoting plant growth and defense against biotic and abiotic stress. However, very little is known about the microbial endophytes living in the spermosphere. Here, we isolated bacteria from the seeds of five different populations of wild cabbage (Brassica oleracea L) that grow within 15 km of each other along the Dorset coast in the UK. The seeds of each plant population contained a unique microbiome. Sequencing of the 16S rRNA genes revealed that these bacteria belong to three different phyla (Actinobacteria, Firmicutes, and Proteobacteria). Isolated endophytic bacteria were grown in monocultures or mixtures and the effects of bacterial volatile organic compounds (VOCs) on the growth and development on B. oleracea and on resistance against a insect herbivore was evaluated. Our results reveal that the VOCs emitted by the endophytic bacteria had a profound effect on plant development but only a minor effect on resistance against an herbivore of B. oleracea. Plants exposed to bacterial VOCs showed faster seed germination and seedling development. Furthermore, seed endophytic bacteria exhibited activity via volatiles against the plant pathogen F. culmorum. Hence, our results illustrate the ecological importance of the bacterial seed microbiome for host plant health and development.

Exploiting chemical ecology to manage hyperparasitoids in biological control of arthropod pests.

Insect hyperparasitoids are fourth trophic level organisms that commonly occur in terrestrial food webs, yet they are relatively understudied. These top-carnivores can disrupt biological pest control by suppressing the populations of their parasitoid hosts, leading to pest outbreaks, especially in confined environments such as greenhouses where augmentative biological control is used. There is no effective eco-friendly strategy that can be used to control hyperparasitoids. Recent advances in the chemical ecology of hyperparasitoid foraging behavior have opened opportunities for manipulating these top-carnivores in such a way that biological pest control becomes more efficient. We propose various infochemical-based strategies to manage hyperparasitoids. We suggest that a push-pull strategy could be a promising approach to 'push' hyperparasitoids away from their parasitoid hosts and 'pull' them into traps. Additionally, we discuss how infochemicals can be used to develop innovative tools improving biological pest control (i) to restrict accessibility of resources (e.g. sugars and alternative hosts) to primary parasitoid only or (ii) to monitor hyperparasitoid presence in the crop for early detection. We also identify important missing information in order to control hyperparasitoids and outline what research is needed to reach this goal. Testing the efficacy of synthetic infochemicals in confined environments is a crucial step towards the implementation of chemical ecology-based approaches targeting hyperparasitoids. © 2019 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Ramanujan's influence on string theory, black holes and moonshine.

Ramanujan influenced many areas of mathematics, but his work on q-series, on the growth of coefficients of modular forms and on mock modular forms stands out for its depth and breadth of applications. I will give a brief overview of how this part of Ramanujan's work has influenced physics with an emphasis on applications to string theory, counting of black hole states and moonshine. This paper contains the material from my presentation at the meeting celebrating the centenary of Ramanujan's election as FRS and adds some additional material on black hole entropy and the AdS/CFT correspondence. This article is part of a discussion meeting issue 'Srinivasa Ramanujan: in celebration of the centenary of his election as FRS'.

Generalism in Nature The Great Misnomer: Aphids and Wasp Parasitoids as Examples.

In the present article we discuss why, in our view, the term 'generalism' to define the dietary breadth of a species is a misnomer and should be revised by entomologists/ecologists with the more exact title relating to the animal in question's level of phagy-mono-, oligo, or polyphagy. We discard generalism as a concept because of the indisputable fact that all living organisms fill a unique ecological niche, and that entry and exit from such niches are the acknowledged routes and mechanisms driving ecological divergence and ultimately speciation. The term specialist is probably still useful and we support its continuing usage simply because all species and lower levels of evolutionary diverge are indeed specialists to a large degree. Using aphids and parasitoid wasps as examples, we provide evidence from the literature that even some apparently highly polyphagous agricultural aphid pest species and their wasp parasitoids are probably not as polyphagous as formerly assumed. We suggest that the shifting of plant hosts by herbivorous insects like aphids, whilst having positive benefits in reducing competition, and reducing antagonists by moving the target organism into 'enemy free space', produces trade-offs in survival, involving relaxed selection in the case of the manicured agro-ecosystem.

Variation in Performance and Resistance to Parasitism of Plutella xylostella Populations.

Two major ecological factors determine the fitness of an insect herbivore: the ability to overcome plant resistance strategies (bottom-up effects) and the ability to avoid or resist attack by natural enemies such as predators and parasitoids (top-down effects). In response to differences in selection pressure, variation may exist in host-plant adaptation and immunity against parasitism among populations of an insect herbivore. We investigated the variation in larval performance of six different Plutella xylostella populations originating from four continents when feeding on a native Dutch plant species, Brassica rapa. One of the used populations has successfully switched its host plant, and is now adapted to pea. In addition, we determined the resistance to attack by the endoparasitoid Diadegma semiclausum originating from the Netherlands (where it is also native) and measured parasitoid performance as a proxy for host resistance against parasitism. Pupal mortality, immature development times, and adult biomass of P. xylostella differed significantly across populations when feeding on the same host plant species. In addition, parasitism success differed in terms of parasitoid adult emergence and their biomass, but not their development times. Variation among natural populations of insects should be considered more when studying interactions between plants and insects up the food chain.