The energetic costs of stereotyped behavior in the paper wasp, Polistes dominulus

Polistes wasps engage in many behavioral interactions. Although there has been debate over the meaning of these interactions, these stereotypical behaviors can be used to determine a colony’s linear dominance hierarchy. Due to the implicit relationship between behavioral and reproductive dominance, behavioral interactions are commonly used to distinguish the reproductively dominant alpha foundress from the beta foundress. It has been suggested that in order to maintain reproductive control, the alpha foundress is forced to remain at a physiologically constrained activity limit. This, in turn, may allow aggressive interactions to be used as determinants influencing reproductive partitioning between cooperating individuals. Energetic costs can place important limitations on behavior, but the energetic cost of the interactions has not previously been measured. To address this, we measured the CO₂ production of 19 non-nestmate pairs displaying interactive and noninteractive behavior. The rate of energy used during interaction behavior was positively associated with published rankings of aggression. However, our results indicate that interactions are not very energetically costly in Polistes, particularly when compared to the likely cost of foraging. These data suggest that maintaining reproductive dominance is not very energetically expensive for the dominant and that the dominant foundress expends energy at a lower rate than the subordinate foundress. Electronic Supplementary Material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Gonadotropic effects of dopamine in isolated workers of the primitively eusocial wasp, Polistes chinensis

In social insects, biogenic amines are thought to play regulatory roles in the transition between reproductive states in females. To determine the effect of dopamine on the reproductive development of workers in primitively eusocial societies, isolated workers of the paper wasp Polistes chinensis were supplied with oral dopamine. Ovarian development was accelerated in dopamine-fed workers as compared to control workers of the same age fed only sucrose solution. Oral dopamine increased brain levels of dopamine and its metabolite (N-acetyldopamine). Brain levels of tyramine or octopamine were also increased by dopamine application in one of two colonies; levels of the tyramine metabolite N-acetyltyramine were unchanged. These results indicate that dopamine plays a gonadotropic role in isolated workers in the primitively eusocial wasp, similar to the gonadotropic role previously reported for juvenile hormone. This is the first study to report effects of dopamine on ovarian development in workers of the paper wasp.     

Examining the “evolution of increased competitive ability” hypothesis in response to parasites and pathogens in the invasive paper wasp Polistes dominula

Successful invaders often become established in new ranges by outcompeting native species. The “evolution of increased competitive ability” hypothesis predicts that invasive species are subjected to less predation and parasitization than sympatric native species, and thus can allocate resources from defence and immunity to growth and fecundity, thereby achieving higher fitness. In this study, we examined whether American invasive Polistes dominula paper wasps have reduced immunocompetence. To explore this scenario, we tested their susceptibility towards parasites and pathogens at both the individual (immune defence) and colony levels, i.e. hygienic behaviour (removal of diseased individuals by nestmates). First, we examined the response to the specific coevolved parasite Xenos vesparum (lost after invasion) in terms of individual host susceptibility and hygienic behaviour. Second, we explored the response against general pathogens by quantifying the bacterial clearance in individual wasps after a challenge with Escherichia coli and hygienic behaviour after a challenge with the fungus Beauveria bassiana. Our results show that American invasive P. dominula have a higher response against X. vesparum at the colony level, but at the individual level their susceptibility is not significantly different from conspecifics of the native range. On the other hand, invasive P. dominula display lower response after a challenge with general pathogens at both the individual and colony levels. While supporting the hypothesis of a reduction of immunocompetence towards general pathogens in invasive species, these findings also suggest that the response against coevolved parasites might follow different evolutionary pathways which are not always easily predictable.     

Absence of within-colony kin discrimination: foundresses of the social wasp, Polistes carolina, do not prefer their own larvae

 There is great potential for conflict within social insect colonies especially when there are multiple inseminated females laying eggs. One reason that conflict is not always realized may be that these females do not identify their own progeny and direct their attentions preferentially towards them. Using DNA microsatellite loci we were able to determine exactly which female was the mother of each larva in eight nests of the social wasp, Polistes carolina. Using 26 h of videotapes of natural nests we observed 2,093 feedings of specific larvae by these adults and found that they did not preferentially feed their own progeny. Instead feedings were distributed to progeny as predicted based on their frequency in the nest. The absence of nepotism towards closest kin within colonies in this system is likely to promote colony harmony. Received: 24 January 2000 / Accepted in revised form: 30 March 2000     

Caste-fate determination primarily occurs after adult emergence in a primitively eusocial paper wasp: significance of the photoperiod during the adult stage

Independent-founding paper wasps constitute a major group of primitively eusocial insects, and when caste-fate determination occurs in temperate species of these wasps, particularly regarding whether it occurs before or after emergence, remains unclear. No critical morphological differences occur between potential queens of the next generation (often called gynes) and workers in primitively eusocial insects. The gynes of temperate species are characterized by diapausing, and the nutrients available during the larval stage have often been believed to determine caste fate. Short days usually induce diapause in temperate nonsocial insects, although few investigations of the effects of day length on caste-fate determination in paper wasps have been conducted. By exposing individuals to different combinations of short and long days during the immature and adult stages, we show for the first time that short days during the adult stage (but not during the immature stage) facilitated caste-fate determination toward gynes in a paper wasp. Moreover, the decision to diapause partly depended on changes in the photoperiod during the pupal and adult stages. The size of the adult also affected caste-fate determination, with diapause more likely to occur in large adults, but this size effect did not occur when individuals were exposed to many short days during the pupal stage. In addition, all adults except for a small proportion of smaller individuals prepared for diapause under short days. These findings suggest that the photoperiod is a higher priority cue than adult size.     

The cues have it; nest-based, cue-mediated recruitment to carbohydrate resources in a swarm-founding social wasp

This study explores whether or not foragers of the Neotropical swarm-founding wasp Polybia occidentalis use nest-based recruitment to direct colony mates to carbohydrate resources. Recruitment allows social insect colonies to rapidly exploit ephemeral resources, an ability especially advantageous to species such as P. occidentalis, which store nectar and prey in their nests. Although recruitment is often defined as being strictly signal mediated, it can also occur via cue-mediated information transfer. Previous studies indicated that P. occidentalis employs local enhancement, a type of cue-mediated recruitment in which the presence of conspecifics at a site attracts foragers. This recruitment is resource-based, and as such, is a blunt recruitment tool, which does not exclude non-colony mates. We therefore investigated whether P. occidentalis also employs a form of nest-based recruitment. A scented sucrose solution was applied directly to the nest. This mimicked a scented carbohydrate resource brought back by employed foragers, but, as foragers were not allowed to return to the nest with the resource, there was no possibility for on-nest recruitment behavior. Foragers were offered two dishes—one containing the test scent and the other an alternate scent. Foragers chose the test scent more often, signifying that its presence in the nest induces naïve foragers to search for it off-nest. P. occidentalis, therefore, employs a form of nest-based recruitment to carbohydrate resources that is mediated by a cue, the presence of a scented resource in the nest.     

Nest odor dynamics in the social wasp Vespula vulgaris

We investigated nest odor dynamics in the common yellow jacket, Vespula vulgaris. In six isolated colonies, we tested the aggression rates toward dead nestmates that had been stored for 10 min, 10 and 19 days outside their colonies at –76 °C. The aggression rate increased from about 12% toward recently killed nestmates up to 30% toward nestmates killed 19 days before the experiment. Obviously, the conserved nest odor profile of the nestmates frozen for several days did not match with that of their colony anymore. This indicates a change of the nest odor within the colony. In a second experiment, we kept two colonies each in one nest box with a complete separation of both neighbor nests by a solid wall inside the box for 28 days. In confrontation experiments, the colony members treated dead foragers from the neighbor nest as aggressively as dead foreign, non-neighbor workers (about 39% each) whereas only about 14% reacted aggressively toward dead nestmates. Seventeen days after the replacement of the solid wall by a metallic grid, which allowed no physical contact but air exchange between the two neighbor colonies, the aggression rates toward foreign workers and nestmates remained relatively unaffected whereas it decreased significantly toward dead neighbors to about 11%. These results suggest a nest odor dynamic caused by volatiles transferred between two adjacent colonies, resulting in an equalization of the former colony specific nest odors. A change of nest odor dynamics influenced by volatiles was so far described only for one ant species at all.     

Two experimental tests of the relationship between group stability and aggressive conflict in Polistes wasps

Two field experiments were used to examine how the relative benefits of cooperation influence within-group conflict in foundress associations of the paper wasp Polistes dominulus. P. dominulus foundresses can either nest alone or cooperate with other foundresses. We experimentally manipulated the relative benefits of co-foundress associations vs independent reproduction and tested the effect on aggressive within-group conflict. First, we examined aggression between alpha and beta co-foundresses before and after lower-ranking foundresses were removed. Removal of subordinates increases the relative contributions of the remaining subordinates to group reproductive output as there are fewer adults to care for the brood. Transactional models predict that group conflict over reproductive shares will increase as the relative benefits of grouping increase. As predicted, aggression between the co-foundresses significantly increased following subordinate removal. Second, we experimentally reduced ecological constraints on independent nesting by placing a previously orphaned, adoptable nest comb near the occupied nests. Providing an independent breeding opportunity is predicted to increase the benefits of independent reproduction relative to those of cooperating, thereby reducing group stability and aggression. As predicted, aggression between dominant and subordinate foundresses significantly decreased after the orphaned comb was presented. Therefore, group members sense variation in ecological constraints and relative productivity contributions and quickly modulate their behavior in response. Overall, these two experiments suggest that paper wasps behave as if within-group competition is limited by the threat of group dissolution such that stable groups where cooperation is strongly favored can withstand higher levels of conflict than unstable groups.     

The contribution of the paper cycle to global warming

Worldwide, paper production is a major industry that contributes about 3 percent of Gross World Product. The paper cycle involves a broad range of natural resource and environmental impacts because fiber supply relies on trees, paper manufacturing requires fuel inputs, and paper waste disposal can contribute to emissions of the potent greenhouse gas (GHG), methane (CH₄). In some countries, the paper cycle may be seen as a net sink for GHG because of reliance on renewable wood by-products and the maintenance of forest plantations. On a worldwide basis, however, this study demonstrates that the paper cycle is a significant contributor to GHG emissions, adding emissions at least comparable in magnitude to that of Australia each year. The estimated global warming contribution of paper in landfills is estimated to be similar to that of paper manufacturing processes, on a heating-equivalent basis. In some temperate regions, original old-growth forests are still harvested to supply pulpwood, resulting in a significant loss of carbon (C) storage. In theory, the paper cycle holds the promise of achieving zero net emissions if pulpwood production, consumption and disposal are carefully managed. In practice, even stabilization of emissions at current levels would be challenging and entail changes comparable to a 20 percent reduction in CH₄ generation from landfilled paper, and a 2.5 percent annual increase in plantation establishment would be needed to offset the projected increase in emissions from paper manufacturing.     

The ways to prevent pollution of the pulp and paper industry in China

Water pollution from the pulp and paper industry is one of the most serious environment problems in China. In order to prevent and treat pollution of the pulp and paper industry, the following works have been done in recent years: 1. Making necessary policies, measures and regulations; 2. Making overall planning and rational layout; 3. Relying mainly on internal treatment developing external treatment as supplement; 4. Improving the technology of treating wastewater of the pulp and paper industry.     

Behavioral evidence for host fidelity among populations of the parasitic wasp, Diachasma alloeum (Muesebeck)

The concept of host fidelity, where host-specific mating occurs in close proximity to the oviposition site and location of larval development, is thought to impart a pre-mating isolation mechanism for sympatric speciation (sensu members of the genus Rhagoletis). The apple maggot fly, Rhagoletis pomonella, and the blueberry maggot fly, R. mendax, are morphologically similar sibling species thought to have speciated in sympatry by divergence of host plant association. Both of these fly species are attacked by the specialist braconid parasitoid, Diachasma alloeum. The current study demonstrates that both male and female D. alloeum exhibit a behavioral preference for the odor of the fruit of their larval Rhagoletis host species. Specifically, those D. alloeum emerging from puparia of R. pomonella are preferentially attracted to hawthorn fruit and those emerging from puparia of R. mendax are preferentially attracted to blueberry fruit. However, male D. alloeum reared from either R. pomonella or R. mendax were equally attracted to females originating from both Rhagoletis species. We suggest that the data herein present evidence for host fidelity, where populations of D. alloeum exhibit a greater tendency to mate and reproduce among the host plants of their preferred Rhagoletis hosts. Furthermore, host fidelity may have resulted in the evolution of distinct host races of D. alloeum tracking the speciation of their larval Rhagoletis prey.     

The effects of different predator species on antipredator behavior in the Trinidadian guppy, Poecilia reticulata

Different types of predators often elicit different antipredator responses in a common type of prey. Alternatively, some prey species may adopt a general response, which provides limited protection from many different types of predator. The Trinidadian guppy, Poecilia reticulata, is faced with a wide range of different predators throughout its range and is known to display varying levels of antipredator behavior depending on the predator assemblage. Pike cichlids, Crenicichla frenata, are regarded as the primary aquatic guppy predator in streams in the northern mountain range in Trinidad. As such, they are seen to be responsible for many of the differences in morphology, life history traits, and behavior between guppy populations from areas with few predators and those from areas with many pike cichlids. In this study we investigated how guppies responded when faced with different predator species using three common aquatic predators. We exposed shoals of ten guppies to one out of four treatments: no predator (control), pike cichlid, acara cichlid (Aequidens pulcher), and wolf fish (Hoplias malabaricus); and we made behavioral observations on both focal individuals and the shoal as a whole. Guppies showed significantly greater levels of predator inspection and shoaling behavior, foraged less, spent more time in the surface water, and stayed in significantly larger shoals when faced with pike cichlids than in other treatments. We discuss these results in the context of multiple predator effects.     

Prehibernating aggregations of Polistes dominulus: an occasion to study early dominance assessment in social insects

At a very early age several mammals establish a first dominance hierarchy, which often persists into adulthood. Social wasps offer an excellent opportunity to study such a phenomenon in insects. Indeed, foundresses of several paper wasps meet in clusters to hibernate from September to March. In spring, wasps found new associative nests where linear hierarchies occur. In the first phase of hibernation, clustering Polistes dominulus wasps show most of the social interactions occurring on the nest 6 months later. At the emergence from diapause, some females already show some behavioral and physiological traits typical of dominant individuals. Here, we investigated the significance of the interactions in the autumnal clusters. We demonstrated that in a given pair, it is more likely that the dominant wasp in autumn becomes the alpha female in spring after the nest foundation phase occurred. Moreover, we showed that dominant females in clusters have both larger body size and ovaries. As ovarian development mainly depends on the social context, our findings seem to indicate that social factors affect the tendency to dominate in aggregations. Furthermore, we suggest that some females may reinforce their physiological status by dominating in clusters, thus increasing the probability to become dominant in spring.     

造纸废水曝气池中钙的去除及迁移转化规律研究

针对造纸废水好氧处理中钙离子（Ca²⁺）浓度过高影响生化过程和后续膜处理的问题,在实验室模拟废水曝气,研究Ca²⁺存在对COD去除效果的影响;对厌氧出水进行曝气实验,探究曝气量对钙去除效果的影响;分析表征造纸废水液相中Ca²⁺浓度和固相污泥中钙的形态,探究造纸废水曝气池中钙的迁移转化规律.结果发现：Ca²⁺的存在会影响活性污泥的活性,进而导致曝气过程中COD的去除率从71.76%降至36.94%;厌氧出水曝气初始阶段内曝气量越大,Ca²⁺的去除率越高;Ca²⁺浓度从厌氧出水到二沉池出水过程中逐渐下降,从游离态转变为CaCO₃的结合态而除去.