Signaling hunger through aggression—the regulation of foraging in a primitively eusocial wasp

Primitively eusocial wasps are generally headed by behaviorally dominant queens who use their aggression to suppress worker reproduction. In contrast, queens in the primitively eusocial wasp Ropalidia marginata are strikingly docile and non-aggressive. However, workers exhibit dominance-subordinate interactions among themselves. These interactions do not appear to reflect reproductive competition because there is no correlation between the relative position of an individual in the dominance hierarchy of the colony and the likelihood that she will succeed a lost/removed queen. Based on the observation that foraging continues unaltered in the absence of the queen and the correlation between dominance behavior and foraging, we have previously suggested that dominance-subordinate interactions among workers in R. marginata have been co-opted to serve the function of decentralized, self-organized regulation of foraging. This idea has been supported by an earlier experimental study where it was found that a reduced demand for food led to a significant decrease in dominance behavior. In this study, we perform the converse experiment, demonstrate that dominance behavior increases under conditions of starvation, and thus provide further evidence in support of the hypothesis that intranidal workers signal hunger through aggression.     

Of Domestic and Wild Guinea Pigs: Studies in Sociophysiology, Domestication, and Social Evolution

Cavia aperea   and Galea musteloides) and domestic (Cavia aperea f. porcellus) guinea pigs, which summarizes general rules for many group-living mammals. It is shown that social interactions have consequences not only for the individuals' reproductive success but also for their degrees of stress and welfare. The way in which individuals interact is controlled not only by the present environment but also by the previous social experiences which they have gathered during their behavioral development. Furthermore, the study of ontogeny does not begin at birth, because prenatal social factors acting on pregnant females can also affect the way in which the offspring will interact when adult. In addition, to understand the genesis of interactions between domesticated animals implies knowledge of the behavioral and physiological changes which occurred during the process of domestication. Finally, understanding the social interactions among individuals of the wild ancestor of the domesticated form requires knowledge of how their behavior patterns were brought about by natural selection during the process of social evolution.     

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.     

The function of dart behavior in the paper wasp, Polistes fuscatus

Dominance behavior in Polistes wasps is a composite trait consisting of various discrete behaviors such as darts, lunges, bites, and mounts. The majority of these behaviors are considered aggressive, and these aggressive behaviors are considered to form a continuum from mild (e.g., darts) to severe (e.g., falling fights). In this paper we focus on darts, the most common of the dominance behaviors, and investigate their function in un-manipulated post-emergent colonies of the primitively eusocial wasp P. fuscatus. Here we show that darts are correlated with the more severe dominance behaviors, and that dominance ranks do not change with the addition or exclusion of darts. We find no correlation, however, between receiving darts and receiving more severe dominance behaviors. This result suggests that darts are not indicative of aggressive reinforcement of dominance, but rather may serve a different function. Our data suggest that the function of darts is to regulate activity on nests. Both foundresses and workers dart inactive workers significantly more often than by chance, and workers respond to a foundresss (but not a workers) dart by becoming less inactive. We also found that active workers who receive a dart from either a foundress or worker respond mostly by switching from one activity to another. Thus, our data suggest that darts are not aggressive behaviors, that foundresses use this signal to initiate activity, and that foundresses and workers both use the signal to regulate worker activity.     

Behavioral genomics of honeybee foraging and nest defense

The honeybee has been the most important insect species for study of social behavior. The recently released draft genomic sequence for the bee will accelerate honeybee behavioral genetics. Although we lack sufficient tools to manipulate this genome easily, quantitative trait loci (QTLs) that influence natural variation in behavior have been identified and tested for their effects on correlated behavioral traits. We review what is known about the genetics and physiology of two behavioral traits in honeybees, foraging specialization (pollen versus nectar), and defensive behavior, and present evidence that map-based cloning of genes is more feasible in the bee than in other metazoans. We also present bioinformatic analyses of candidate genes within QTL confidence intervals (CIs). The high recombination rate of the bee made it possible to narrow the search to regions containing only 17–61 predicted peptides for each QTL, although CIs covered large genetic distances. Knowledge of correlated behavioral traits, comparative bioinformatics, and expression assays facilitated evaluation of candidate genes. An overrepresentation of genes involved in ovarian development and insulin-like signaling components within pollen foraging QTL regions suggests that an ancestral reproductive gene network was co-opted during the evolution of foraging specialization. The major QTL influencing defensive/aggressive behavior contains orthologs of genes involved in central nervous system activity and neurogenesis. Candidates at the other two defensive-behavior QTLs include modulators of sensory signaling (Am5HT ₇ serotonin receptor, AmArr4 arrestin, and GABA-B-R1 receptor). These studies are the first step in linking natural variation in honeybee social behavior to the identification of underlying genes.     

Indirect interactions between invasive and native plants via pollinators

In generalised pollination systems, the presence of alien plant species may change the foraging behaviour of pollinators on native plant species, which could result in reduced reproductive success of native plant species. We tested this idea of indirect interactions on a small spatial and temporal scale in a field study in Mauritius, where the invasive strawberry guava, Psidium cattleianum, provides additional floral resources for insect pollinators. We predicted that the presence of flowering guava would indirectly and negatively affect the reproductive success of the endemic plant Bertiera zaluzania, which has similar flowers, by diverting shared pollinators. We removed P. cattleianum flowers within a 5-m radius from around half the B. zaluzania target plants (treatment) and left P. cattleianum flowers intact around the other half (control). By far, the most abundant and shared pollinator was the introduced honey bee, Apis mellifera, but its visitation rates to treatment and control plants were similar. Likewise, fruit and seed set and fruit size and weight of B. zaluzania were not influenced by the presence of P. cattleianum flowers. Although other studies have shown small-scale effects of alien plant species on neighbouring natives, we found no evidence for such negative indirect interactions in our system. The dominance of introduced, established A. mellifera indicates their replacement of native insect flower visitors and their function as pollinators of native plant species. However, the pollination effectiveness of A. mellifera in comparison to native pollinators is unknown. Christine B. Müller, deceased 7 March 2008.     

RNAi-mediated silencing of vitellogenin gene function turns honeybee (Apis mellifera) workers into extremely precocious foragers

The switch from within-hive activities to foraging behavior is a major transition in the life cycle of a honeybee (Apis mellifera) worker. A prominent regulatory role in this switch has long been attributed to juvenile hormone (JH), but recent evidence also points to the yolk precursor protein vitellogenin as a major player in behavioral development. In the present study, we injected vitellogenin double-stranded RNA (dsVg) into newly emerged worker bees of Africanized genetic origin and introduced them together with controls into observation hives to record flight behavior. RNA interference-mediated silencing of vitellogenin gene function shifted the onset of long-duration flights (>10 min) to earlier in life (by 3–4 days) when compared with sham and untreated control bees. In fact, dsVg bees were observed conducting such flights extremely precociously, when only 3 days old. Short-duration flights (<10 min), which bees usually perform for orientation and cleaning, were not affected. Additionally, we found that the JH titer in dsVg bees collected after 7 days was not significantly different from the controls. The finding that depletion of the vitellogenin titer can drive young bees to become extremely precocious foragers could imply that vitellogenin is the primary switch signal. At this young age, downregulation of vitellogenin gene activity apparently had little effect on the JH titer. As this unexpected finding stands in contrast with previous results on the vitellogenin/JH interaction at a later age, when bees normally become foragers, we propose a three-step sequence in the constellation of physiological parameters underlying behavioral development. David Santos Marco Antonio and Karina Rosa Guidugli-Lazzarini contributed equally to the present study.     

Suppression pheromone and cockroach rank formation

Although agonistic behaviors in the male lobster cockroach (Nauphoeta cinerea) are well known, the formation of an unstable hierarchy has long been a puzzle. In this study, we investigate how the unstable dominance hierarchy in N. cinerea is maintained via a pheromone signaling system. In agonistic interactions, aggressive posture (AP) is an important behavioral index of aggression. This study showed that, during the formation of a governing hierarchy, thousands of nanograms of 3-hydroxy-2-butanone (3H-2B) were released by the AP-adopting dominant in the first encounter fight, then during the early domination period and that this release of 3H-2B was related to rank maintenance, but not to rank establishment. For rank maintenance, 3H-2B functioned as a suppression pheromone, which suppressed the fighting capability of rivals and kept them in a submissive state. During the period of rank maintenance, as the dominant male gradually decreased his 3H-2B release, the fighting ability of the subordinate gradually developed, as shown by the increasing odds of a subordinate adopting an AP (OSAP). The OSAP was negatively correlated with the amount of 3H-2B released by the dominant and positively correlated with the number of domination days. The same OSAP could be achieved earlier by reducing the amount of 3H-2B released by the dominant indicates that whether the subordinate adopts an offensive strategy depends on what the dominant is doing.     

Costs and benefits of subordinate queens in colonies of the ant,<Emphasis Type="Italic"> Leptothorax gredleri</Emphasis>

Queens of some species of social insects form dominance hierarchies in which only the top-ranking individual reproduces. Such unequal partitioning of reproduction can be stable when subordinate queens increase their inclusive fitness indirectly by helping to rear a related dominant queen's offspring. We investigated whether subordinate queens of the ant, Leptothorax gredleri, affect the reproductive success of the dominant queen. In laboratory colonies with two queens, first eggs were laid approximately 20 days later than in colonies with a single queen, presumably due to aggressive competition among nestmate queens. Nevertheless, two-queen colonies produced on average more brood than one-queen colonies, although egg laying was completely or almost completely monopolized by one single queen. The presence of a subordinate queen therefore appears to increase the productivity of the dominant.     

Strategic 3-hydroxy-2-butanone release in the dominant male lobster cockroach, <Emphasis Type="Italic">Nauphoeta cinerea</Emphasis>

In the lobster cockroach Nauphoete cinerea, the dominant–subordinate hierarchy formed via the agonistic interactions is unstable, and changes in rank order are common. Our previous results showed that in the first encounter fight during initial rank formation, microgram levels of 3H-2B are released by the aggressive posture (AP)-adopting dominant male. In the present study, the pattern of daily pheromone (3H-2B) release during the domination period and on the day of rank switch, rank duration, and rank switch frequency were investigated in three-male groups and six-male groups to examine the effect of higher frequency of agonistic encounters. The results showed that, in the three-male groups (50-day observation period), daily 3H-2B release rate was not constant, but fluctuated, the average duration of dominant rank was 16.6 ± 2.0 days, rank switch occurred in 58.8% of groups, and the frequency of rank switching (average number of rank switches/group/50 days) was 1.4 ± 0.2. For the six-male groups (30-day observation period), the daily 3H-2B release rate also fluctuated, but the duration of dominant rank was significantly shorter at 4.2 ± 0.6 days, rank switch occurred in 100% of groups, and the frequency of rank switching (average number of rank switches/group/30 days) was significantly higher at 6.9 ± 0.6. The results for both sets of male groups showed that as a new rank formed (either on the first encounter day or on the day of rank switching), the dominant status was significantly associated with a higher 3H-2B release rate. In the animal kingdom, fighting usually involves communication or the exchange of signals, and the results of this study indicated that the fluctuating daily 3H-2B release rate adopted by the dominants is a kind of strategic release and the 3H-2B release rate is a signal used to determine dominance.     

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.     

2种有机磷农药联合胁迫下日本青鳉的逐级行为响应

采用水质安全在线生物预警系统(BEWs)记录行为强度数据,探讨了2种有机磷农药敌百虫和对硫磷联合胁迫下,不同暴露浓度、不同暴露时间日本青鳉的行为响应.在不同浓度比例的农药暴露中,20、10、5、1和0.1TU的暴露浓度下行为反应时间分别为:1.4、4.8、6.4、32.2和43.0h,并且每个浓度不同配比之间行为反应时间差异不明显.结果表明,在2种有机磷农药的暴露下,日本青鳉的逐级行为响应既受化合物浓度高低的影响,又受暴露时间的影响,但每个浓度的不同配比之间的行为响应时间差异不明显,而且每个浓度2种药物不同配比暴露下的青鳉鱼的逐级行为响应基本一致,即2种作用机制相似的有机磷农药对日本青鳉的行为毒性是简单的相加作用.     

The role of tyramine and octopamine in the regulation of reproduction in queenless worker honeybees

In honeybees, workers under queenless condition compete for reproduction and establish reproductive dominance hierarchy. Ovary activation is generally accompanied by the expression of queen-like pheromones. Biogenic amines (BAs), in particular dopamine, are believed to be involved in this process by regulating ovarian development. However, the role of BAs in establishing reproductive dominance or their effect on queen-like pheromone production was not investigated. Here, we explored the effect of octopamine (OA) and tyramine (TA) oral treatments on the propensity of treated bees to become reproductively dominant and produce queen-like pheromones in Dufour’s and mandibular glands. One bee in a pair was treated with either OA or TA while the other was fed sugar solution. TA was found to enhance ovary development and the production of esters in the Dufour’s gland and 9HDA (queen component) in the mandibular glands, thus facilitating worker reproductive dominance. OA, on the other hand, did not enhance ovarian development or ester production, but increased the production of 10HDA (worker major component) in the mandibular glands of their sugar-paired mates. OA is known to induce foraging behavior by workers, while increased production of 10HDA characterizes nursing workers. Therefore, we suggest that TA induces reproductive division of labor, while OA treatment results in caste differentiation of workers to foragers and nurses.     

The reproductive choices of eavesdropping female black-capped chickadees,<Emphasis Type="Italic"> Poecile atricapillus</Emphasis>

In animals where males engage in signalling interactions, females might evaluate male–male contests to inform their reproductive choices. We used interactive playback to engage territorial male black-capped chickadees (Poecile atricapillus) of known dominance status in countersinging contests with an aggressive or submissive opponent. Previous analysis of these data showed that high-ranking males who received aggressive playback were more likely to be cuckolded. Here we describe the particular reproductive decisions of females whose partners received aggressive versus submissive playback. The proportion of extra-pair young per brood was higher for females paired to high-ranking males that received aggressive playback compared to submissive playback, and similar to levels in broods of females paired to low-ranking males. We found no strong predictors of whether high-ranking subjects lost paternity following aggressive playback. Females usually preferred extra-pair sires with high dominance status. When females had extra-pair fertilizations with low-ranking males, females chose males who had received submissive playback. We conclude that females mated to aggressive-playback, high-ranking males pursued mixed mating strategies similar to those of females mated to low-ranking males. Our results support the idea that male performance in song contests may influence multiple aspects of female reproductive choices.     

不同营养环境下棕鞭藻共栖细菌的群落结构比较

微藻富含油脂,可以用作合成生物柴油的原料,被认为是最具发展前景的生物质能源.微藻的光合培养体系往往是非纯培养体系,现有研究更多地关注微藻生物量的积累及其对废水环境的净化效果,而对体系中广泛存在的共栖细菌缺乏全面深入的认知.本文通过对棕鞭藻共栖细菌的16S rDNA基因进行高通量测序分析,研究棕鞭藻（Ochromonas）在生活废水、BG11及Glu+BG11（BG11中添加10 g·L^-1葡萄糖）3种不同营养环境中共栖细菌的群落结构差异,进而阐明有机物及复杂废水环境对微藻共栖细菌群落结构的影响效果.结果表明,生活废水、BG11及Glu+BG11 3种营养环境中共栖细菌群落结构存在显著差异,生活废水体系中生物多样性显著高于BG11及Glu+BG11体系,生活废水中共栖细菌香浓（Shannon）多样性指数高达4.32,其次是BG11及Glu+BG11,Shannon指数分别为2.13、1.54.从共栖细菌群落的组成上看,3种营养环境中优势菌属差异显著,生活废水中优势菌属有芽殖杆菌属（Gemmobacter）、鞘脂单胞菌属（Sphingomonas）及玫瑰球菌属（Roseococcus）,相对丰度分别为14.46%、14.9%、14.54%,其中,芽殖杆菌属只在生活废水中有较高丰度.BG11中寡养单胞菌属（Stenotrophomonas）与玫瑰球菌属的丰度分别高达26.86%、51.03%.Glu+BG11中寡养单胞菌属较BG11中显著增加,达到82.41%,而玫瑰球菌属较生活废水及BG11中则明显降低,菌群丰度仅为6.2%.对比3种营养环境,玫瑰球菌属均具有较高丰度,是棕鞭藻良好的共生菌.     

Fertility signaling—the proximate mechanism of worker policing in a clonal ant

In eusocial insects, the ability to regulate reproduction relies on cues that signal the presence of fertile individuals. We investigated the variation of cuticular hydrocarbons (CHCs) with reproductive status in Platythyrea punctata, an ant, in which all workers are capable of producing daughters from unfertilized eggs (thelytoky). Who reproduces is determined through dominance and worker policing. New reproductives, which developed their ovaries after separation from an old reproductive for a short period of time, were attacked by nonreproductives upon reintroduction into their colony. In contrast, aggression against new reproductives with fully developed ovaries, which had been separated over a longer period, was initiated by fights between old and new reproductives. CHC profiles varied with ovarian development. New reproductives were only attacked when they expressed a CHC profile similar to old reproductives, but not when it was similar to that of nonreproductives. CHCs appear to signal the fertility of individuals and induce policing behavior towards surplus reproductive workers.     

Vegetation dynamics after compost amendment in a Mediterranean post-fire ecosystem

On Mediterranean calcareous soils, high fire frequency induces soil impoverishment and the development of stable Quercus coccifera garrigues. Organic amendment could increase soil fertility levels, and could alter the vegetation dynamics and the established dominance relationships. In this study, the plant cover evolution is monitored in an amended burnt shrub during two years. Three treatments are studied: control (D0), 50 t ha⁻¹ (D50) and 100 t ha⁻¹ (D100) of fresh co-composted sewage sludge and greenwastes.First, the spreading process leads to the squashing of the vegetation, whose intensity reaches a threshold on D100 and limits the woody species’ recovery after amendment. Consequently, the dominant herb Brachypodium retusum is favoured. On D50, woody species are favoured compared to herbs, probably due to the space colonization strategy of Q. coccifera after squashing. Thus, compost at both rates favours the two dominant resprouter species until they have colonized all the free space created by squashing.Secondly, compost has some fertilizing effects on Rosmarinus officinalis and Ulex parviflorus, that improves their cover on D50. However, compost also increases U. parviflorus’ sensitivity to drought.At the end of the experiment, a third compost effect appears, as seeder cover becomes greater than D0 on D50. After amendment, compost covers rocky stones, which creates a new territory where species with superficial rooting can establish (mulch effect).     

Sex mosaics in a male dimorphic ant Cardiocondyla kagutsuchi

Gynandromorphy, or the development of organisms with a combination of male and female morphological features, is common in Hymenoptera. The underlying mechanism is likely associated with the sex-determination system, and studying this phenomenon should lead to a deeper understanding of both embryonic development and sex determination. The reproductive capabilities of gynandromorphs (hereafter, sex mosaics) remain unclear. We studied gynandromorphy in the Malaysian ant Cardiocondyla kagutsuchi, which has sex mosaics of queens (gynandromorphs; mosaic of queens and winged male) and workers (ergatandromorphs; mosaic of worker and wingless ergatoid male). These sex mosaics were classified into seven morphological categories. Most individuals had more male than female body areas. Behavioral observations revealed that sex mosaics behave more in accordance with the “sex” of their brain than that of the reproductive organs (gaster). Relative DNA quantities showed that both female and male regions contained haploid and diploid nuclei, irrespective of their phenotypic appearance, indicating that external appearance did not reflect internal tissues. Nearly one third of the adults were sex mosaics and they were not infected with Wolbachia. Our results suggest that the production of sex mosaics in this species does not pose a substantial cost to colonies and that the underlying causes are therefore not strongly selected against.     

Rapid modification in the olfactory signal of ants following a change in reproductive status

In insect societies, the presence and condition of egg-layers can be assessed with pheromones. Exocrine secretions are expected to vary in time in order to give up-to-date information on an individuals reproductive physiology. In the queenless monogynous ant Streblognathus peetersi, we allowed a previously infertile high-ranking worker to accede to the alpha rank, thus triggering the onset of her oogenesis (15 replicates). We then studied her interactions with an established egg-layer from the same colony after different durations, ranging from 20 h to several days. Even though her eggs are only ready to be laid after 30 days, the new alpha was recognised within 1–2 days. Detection occurred at a distance of a few millimetres, suggesting the involvement of a pheromone with low volatility, such as cuticular hydrocarbons. When the new alpha had differentiated for >48 h, she was attacked by the established egg-layer. In all cases, low-ranking workers eventually immobilised one of the two alphas: the new alpha was the target if she had differentiated only recently, suggesting that police workers select the dominant worker with the less fertile odour. Using the behaviour of ants as our measure, we demonstrate that a dominants olfactory signal changes rapidly with a modification in her social status, and it occurs well before the onset of egg-laying.