Felix L. Wäckers

Neonicotinoid pesticide reduces bumble bee colony growth and queen production.

Bad News for Bees Neonicotinoid insecticides were introduced in the early 1990s and have become one of the most widely used crop pesticides in the world. These compounds act on the insect central nervous system, and they have been shown to be persistent in the environment and in plant tissues. Recently, there have been controversial connections made between neonicotinoids and pollinator deaths, but the mechanisms underlying these potential deaths have remained unknown. Whitehorn et al. (p. 351, published online 29 March) exposed developing colonies of bumble bees to low levels of the neonicotinoid imidacloprid and then released them to forage under natural conditions. Treated colonies displayed reduced colony growth and less reproductive success, and they produced significantly fewer queens to found subsequent generations. Henry et al. (p. 348, published online 29 March) documented the effects of low-dose, nonlethal intoxication of another widely used neonicotinoid, thiamethoxam, on wild foraging honey bees. Radio-frequency identification tags were used to determine navigation success of treated foragers, which suggested that their homing success was much reduced relative to untreated foragers. Bumble bee colonies produce many fewer queens after exposure to a widely used insecticide. Growing evidence for declines in bee populations has caused great concern because of the valuable ecosystem services they provide. Neonicotinoid insecticides have been implicated in these declines because they occur at trace levels in the nectar and pollen of crop plants. We exposed colonies of the bumble bee Bombus terrestris in the laboratory to field-realistic levels of the neonicotinoid imidacloprid, then allowed them to develop naturally under field conditions. Treated colonies had a significantly reduced growth rate and suffered an 85% reduction in production of new queens compared with control colonies. Given the scale of use of neonicotinoids, we suggest that they may be having a considerable negative impact on wild bumble bee populations across the developed world.

Linking above- and belowground multitrophic interactions of plants, herbivores, pathogens, and their antagonists

Plants function in a complex multitrophic environment. Most multitrophic studies, however, have almost exclusively focused on aboveground interactions, generally neglecting the fact that above- and belowground organisms interact. The spatial and temporal dynamics of above- and belowground herbivores, plant pathogens, and their antagonists, can differ in space and time. This affects the temporal interaction strengths and impacts of above- and belowground higher trophic level organisms on plants. Combining both above- and belowground compartments in studies of multitrophic interactions throughout the life cycle of plants will improve our understanding of ecology and evolution in the real world.

Learning of Host-Finding Cues by Hymenopterous Parasitoids

Interactions between insect parasitoids and their arthropod hosts characteristically result in the premature death of the hosts, and are obligatory for the development of the parasitic insects. This obviously places strong pressure on the hosts to avoid detection by parasitoids, and on the parasitoids themselves to improve encounter rates with suitable hosts. To confront the challenge of finding the often-inconspicuous, well-hidden hosts, parasitoids have developed various sophisticated searching strategies that depend on a vast array of environmental cues.

Recruitment of predators and parasitoids by herbivore-injured plants

Introduction In recent years, induced plant defenses have received widespread attention from biologists in a variety of disciplines. The mechanisms underlying these defenses and the interactions that mediate them appeal not only to plant physiologists, ecologists, and evolutionary biologists but also to those scientists that search for novel strategies in plant protection. Several recent books (Karban and Baldwin, 1997; Agrawal et al. , 1999) and reviews (Baldwin, 1994; Karban et al. , 1997; Agrawal and Rutter, 1998; Agrawal and Karban, 1999; Baldwin and Preston, 1999; Dicke et al. , 2003) have been devoted entirely to the subject of induced plant defenses. Various forces, ranging from abiotic stresses to biotic factors such as pathogens, arthropods, or higher organisms, may trigger different plant defense responses. Yet, the biochemical pathways that are involved appear to show considerable similarities. This is also true for the so-called indirect defenses. The term indirect defense refers to those adaptations that result in the recruitment and sustenance of organisms that protect the plants against herbivorous attackers. The early published examples of indirect defenses involved intimate plant–ant interactions, in which myrmecophilous plants were shown to have evolved a range of adaptations providing ants with shelter (domatia) and various food sources (Belt, 1874; Janzen, 1966). In return, these plants may obtain a range of benefits because ants can provide nutrition (Thomson, 1981) or more commonly, protection against herbivores, pathogens, and competing plants (e.g. Koptur, 1992; Oliveira, 1997).

Plant-provided food for carnivorous insects : a protective mutualism and its applications

Plants provide insects with a range of specific foods, such as nectar, pollen, and food bodies. In exchange, they may obtain various services from arthropods. The role of food rewards in the plant–pollinator mutualism has been broadly covered. This book addresses another category of food-mediated interactions, focussing on how plants employ foods to recruit arthropod ‘‘bodyguards’’ as a protection against herbivores.

How to hunt for hiding hosts: the reliability-detectability problem in foraging parasitoids.

Foraging parasitoids may use stimuli that are derived from their host or from the food of their host, often plants. But how usable are 2nd and 1st trophic level stimuli and what is their relative importance in parasitoid foraging? It is argued that foraging parasitoids are facing a reliability-detectability problem: host-derived stimuli are the most reliable in indicating host presence, accessibility and suitability but they are generally hard to detect. Plant stimuli, on the other hand, are easier to detect but arc generally less reliable indicators. Parasitoids have evolved different non-exclusive strategies to solve this problem. (1) Infochemical detour: parasitoids resort to information from other, more detectable, host stages than the one under attack. (2) Herbivore-induced synomones: parasitoids use specific plant produced volatiles that are released upon damage by a specific herbivore species. In the present paper we put most emphasis on a third strategy (3) Associative learning: through associative learning parasitoids link easy-to-detect stimuli to reliable but hard-to-detect stimuli. Specific mechanisms by which associative learning can improve foraging success are discussed.

Conserved class of queen pheromones stops social insect workers from reproducing

Long Live the Queen Eusociality is often considered to have arisen, at least in part, due to the inclusive fitness that workers gain through helping their queen sister to raise her offspring. Van Oystaeyen et al. (p. 287; see the Perspective by Chapuisat) characterized the sterility-inducing queen pheromone across three distantly related eusocial hymenopterans (a wasp, a bumblebee, and a desert ant) and synthesized data across 69 other species. Queen pheromones appear to be remarkably conserved, which suggests that reproductive manipulation has ancient roots. Social insect queens use an ancient, evolutionarily conserved class of pheromones to prevent worker reproduction. [Also see Perspective by Chapuisat] A major evolutionary transition to eusociality with reproductive division of labor between queens and workers has arisen independently at least 10 times in the ants, bees, and wasps. Pheromones produced by queens are thought to play a key role in regulating this complex social system, but their evolutionary history remains unknown. Here, we identify the first sterility-inducing queen pheromones in a wasp, bumblebee, and desert ant and synthesize existing data on compounds that characterize female fecundity in 64 species of social insects. Our results show that queen pheromones are strikingly conserved across at least three independent origins of eusociality, with wasps, ants, and some bees all appearing to use nonvolatile, saturated hydrocarbons to advertise fecundity and/or suppress worker reproduction. These results suggest that queen pheromones evolved from conserved signals of solitary ancestors.

High Susceptibility of Bt Maize to Aphids Enhances the Performance of Parasitoids of Lepidopteran Pests

Concerns about possible undesired environmental effects of transgenic crops have prompted numerous evaluations of such crops. So-called Bt crops receive particular attention because they carry bacteria-derived genes coding for insecticidal proteins that might negatively affect non-target arthropods. Here we show a remarkable positive effect of Bt maize on the performance of the corn leaf aphid Rhopalosiphum maidis, which in turn enhanced the performance of parasitic wasps that feed on aphid honeydew. Within five out of six pairs that were evaluated, transgenic maize lines were significantly more susceptible to aphids than their near-isogenic equivalents, with the remaining pair being equally susceptible. The aphids feed from the phloem sieve element content and analyses of this sap in selected maize lines revealed marginally, but significantly higher amino acid levels in Bt maize, which might partially explain the observed increased aphid performance. Larger colony densities of aphids on Bt plants resulted in an increased production of honeydew that can be used as food by beneficial insects. Indeed, Cotesia marginiventris, a parasitoid of lepidopteran pests, lived longer and parasitized more pest caterpillars in the presence of aphid-infested Bt maize than in the presence of aphid-infested isogenic maize. Hence, depending on aphid pest thresholds, the observed increased susceptibility of Bt maize to aphids may be either a welcome or an undesirable side effect.

INDUCIBLE DEFENSES AND TROPHIC STRUCTURE

Resource edibility is a crucial factor in ecological theory on the relative importance of bottom-up and top-down control. Current theory explains trophic structure in terms of the relative abundance and succession of edible and inedible species across gradients of primary productivity. We argue that this explanation is incomplete owing to its focus on inedibility and the assumption that plants and herbivores have fixed defense levels. Consumer-induced defenses are an important source of variation in the vulnerability of prey and are prevalent in natural communities. Such induced defenses decrease per capita consumption rates of consumers but hardly ever result in complete inedibility. When defenses are inducible a prey population may consist of both undefended and defended individuals. Here we use food chain models with realistic parameter values to show that variation in consumption rates on different prey types causes a gradual instead of stepwise increase in the biomass of all trophic levels in response to enrichment. Such all-level responses have been observed in both aquatic and terrestrial ecosystems and in microbial food chains in the laboratory. We stress that, in addition to the known food web effects of interspecific variation in edibility, intraspecific variation in edibility is another form of within-trophic-level heterogeneity that also has such effects. We conclude that inducible defenses increase the relative importance of bottom-up control.

Effects of honeydew sugar composition on the longevity of Aphidius ervi

Feeding on sugar‐rich foods such as nectar and honeydew is important for survival of many adult parasitoids. Especially in agricultural systems, honeydew is often the most prevalent carbohydrate source. However, relative to plant nectar, honeydew may be relatively unsuitable, as a result of an unfavourable sugar composition or the presence of secondary plant compounds. We studied survival of the aphid parasitoid Aphidius ervi Haliday (Hymenoptera: Braconidae) on honeydew collected from various aphid species feeding on potato (Solanum tuberosum L., cv. Desiree) (Solanaceae), wheat (Triticum aestivum L., cv. Bobwhite) (Poaceae), or artificial diet, as well as the sugar composition of the different honeydews. Honeydews from the tested aphid species on potato, wheat, or artificial diet were found to be relatively suitable food sources for adult A. ervi, although not always as suitable as a 2 M sucrose solution. There were differences in honeydew sugar composition among the different aphid species on the various host plants. Multivariate statistics showed that the factor ‘aphid species’ had a significant influence on the sugar composition of the honeydew, explaining 27% of the variation in the potato system and 89% in the wheat system. When exploring the relationship between carbohydrate composition of the honeydews from aphids on potato and wheat plants, and their nutritional value for A. ervi, data revealed that differences in parasitoid longevity can to some extent be explained by carbohydrate composition. Furthermore, our results confirm that sucrose and its hexose components glucose and fructose are very suitable carbohydrate sources for hymenopteran parasitoids and show that parasitoid survival on an equimolar solution of the two monosaccharides glucose and fructose does not exceed performance on the disaccharide sucrose.