Selfish strategies and honest signalling: reproductive conflicts in ant queen associations

Social insects offer unique opportunities to test predictions regarding the evolution of cooperation, life histories and communication. Colony founding by groups of unrelated queens, some of which are later killed, may select for selfish reproductive strategies, honest signalling and punishment. Here, we use a brood transfer experiment to test whether cofounding queens of the ant Lasius niger ‘selfishly’ adjust their productivity when sharing the nest with future competitors. We simultaneously analysed queen cuticular hydrocarbon (CHC) profiles to investigate whether queens honestly signal their reproductive output or produce dishonest, manipulative signals, providing a novel test of the evolutionary significance of queen pheromones. Queens produced fewer workers when their colony contained ample brood, but only in the presence of competitors, suggesting selfish conservation of resources. Several CHCs correlated with reproductive maturation, and to a lesser extent with productivity; the same hydrocarbons were more abundant on queens that were not killed, suggesting that workers select productive queens using these chemical cues. Our results highlight the role of honest signalling in the evolution of cooperation: whenever cheaters can be reliably identified, they may incur sanctions that reduce the incentive to be selfish.     

A social insect fertility signal is dependent on chemical context

Identifying group members and individuals'' status within a group are fundamental tasks in animal societies. For ants, this information is coded in the cuticular hydrocarbon profile. We manipulated profiles of the ant Odontomachus brunneus to examine whether the releaser and primer effects of fertility signals are dependent on chemical context. Fertility status is signalled through increased abundance of (Z)-9-nonacosene (Z9 : C₂₉). Across the ant''s distribution, populations have distinct hydrocarbon profiles but the fertility signal is conserved. Foreign queens and fertility-signal-treated workers from the same population, sharing a similar chemical background, elicited releaser effects from workers, whereas queens and fertility-signal-treated workers from different populations did not. Z9 : C₂₉ presented without chemical background did not elicit releaser effects. A primer-effect experiment found that Z9 : C₂₉, presented without a chemical background, did not inhibit worker reproduction. Our results demonstrate that a familiar chemical background is necessary for appropriate responses to fertility signals.     

Influence of immature queen honeybees (Apis mellifera) on queen rearing and foraging

ABSTRACT. Providing queenless colonies with five queen cells containing larvae or pupae diminished the number of queen cells and queen cell cups subsequently produced, but not as effectively as the provision of a mated laying queen. Immature queens were more effective than the mature queens in stimulating pollen collection, but were less effective in stimulating nectar collection.     

Influence of virgin queen honeybees (Apis mellifera) on queen rearing and foraging

ABSTRACT. Virgin queens are as effective as mated laying queens at inhibiting colonies from rearing queens but not from producing queen cell cups. Colonies without brood produce fewer queen cell cups than similar colonies that have brood. Colonies without queens forage much less and collect less pollen than with either a mated or virgin queen. Colonies with virgin queens forage as much as those with mated queens but collected less pollen.     

Identification of an ant queen pheromone regulating worker sterility

The selective forces that shape and maintain eusocial societies are an enduring puzzle in evolutionary biology. Ordinarily sterile workers can usually reproduce given the right conditions, so the factors regulating reproductive division of labour may provide insight into why eusociality has persisted over evolutionary time. Queen-produced pheromones that affect worker reproduction have been implicated in diverse taxa, including ants, termites, wasps and possibly mole rats, but to date have only been definitively identified in the honeybee. Using the black garden ant Lasius niger, we isolate the first sterility-regulating ant queen pheromone. The pheromone is a cuticular hydrocarbon that comprises the majority of the chemical profile of queens and their eggs, and also affects worker behaviour, by reducing aggression towards objects bearing the pheromone. We further show that the pheromone elicits a strong response in worker antennae and that its production by queens is selectively reduced following an immune challenge. These results suggest that the pheromone has a central role in colony organization and support the hypothesis that worker sterility represents altruistic self-restraint in response to an honest quality signal.     

The role of queen mandibular pheromone and colony congestion in honey bee (Apis mellifera L.) reproductive swarming (Hymenoptera: Apidae)

The roles of honey bee queen mandibular pheromone and colony congestion in the inhibition of swarming were investigated. Two colony siz.es were used: small, congested colonies and large, uncongested colonies. Both groups of colonies were treated with various dosages of the five-component, synthetic queen mandibular pheromone in the spring, and the extent and timing of swarming were followed. Most treatment groups received pheromone or a solvent blank (control) on a stationary slide; one group of the congested colonies received a pheromone treatment via an aerosol spray. The pheromone was not effective at delaying swarming in the congested colonies at any dosage applied on slides, but the aerosol spray-treated colonies swarmed significantly later in the season than the control colonies. The uncongested, pheromone-treated colonies exhibited a dose-dependent delay in swarming, with the highest dosage colonies swarming almost four weeks later than the control colonies. These results indicate an interaction between congestion and pheromone in the control of honey bee reproduction. While congestion may in itself be a factor stimulating swarming, these results are consistent with the interpretation that colony congestion reduces the transmission of queen pheromone within the nest, thereby removing the queen 's pheromone-based inhibition of queen rearing and subsequent swarming by workers.     

Elimination of sexual brood in the Argentine antLinepithema humile: queen effect and brood recognition

Mated queens of the antLinepithema humile (Iridomyrmex humilis Mayr) introduced into dequeened colony fragments rearing sexual brood elicited worker aggression resulting in queen larvae being bitten and eliminated. By contrast, male larvae were spared. Regarding queen brood, killing mainly concerned small and medium sized larvae. A large proportion of the large larvae escaped extermination, and prepupae and pupae were spared. These data suggest that workers were able to discriminate sex, caste and age of the brood. That a queen pheromone may be involved was shown by experiments using whole or cut corpses that were either rinsed or not rinsed in pentane. The pheromone eliciting worker aggressive behaviour was shown to act over a short distance, suggesting that it is somewhat volatile. Similarities and differences between this new queen pheromone and other known queen pheromones acting on queen production or worker attraction are discussed as well as the origin of the signals underlying the recognition of the larval classes.     

Changes in composition of cuticular biochemicals of the facultatively polygynous ant Petalomyrmex phylax during range expansion in Cameroon with respect to social, spatial and genetic variation

In social insects, biochemicals found at the surface of the cuticle are involved in the recognition process and in protection against desiccation and pathogens. However, the relative contribution of evolutionary forces in shaping diversity of these biochemicals remains largely unresolved in ants. We determined the composition of epicuticular biochemicals for workers sampled in 12 populations of the ant Petalomyrmex phylax from Cameroon. Genetic variation at 12 microsatellite markers was used to infer population history and to provide null expectations under the neutrality hypothesis. Genetic data suggest a recent southward range expansion of this ant species. Furthermore, there is a decline southward in the numbers of queens present in mature colonies. Here, we contrast the pattern of biochemical variation against genetic, social and spatial parameters. We thus provide the first estimates of the relative contribution of neutral and selective processes on variation of ant cuticular profile. Populations in migration-drift disequilibrium showed reduction of within-population variation for genetic markers as well as for cuticular profiles. In these populations, the cuticular profile became biased towards a limited number of high molecular weight molecules. Within- and among-population biochemical variation was explained by both genetic and social variation and by the spatial distribution of populations. We therefore propose that during range expansion of P. phylax, the composition of epicuticular compounds has been affected by a combination of neutral processes - genetic drift and spatially limited dispersal - and spatially varying selection, social organization and environmental effects.     

Queen replacement in African and European honey bee colonies with and without afterswarms

We examined the dynamics of the queen replacement process in African and European colonies that did and did not produce afterswarms. In colonies without afterswarms, the queen replacement process was completed in 24–48 hours, the first-emerging virgin queen (VQ) typically inherited the natal nest even if multiple queens emerged, workers performed few vibration signals on emerged queens, and all signaling activity was directed toward early emerging VQs. In contrast, if colonies did produce afterswarms, the queen replacement process required 5–6 days, there was no advantage for first-emerging queens, vibration rates on emerged queens were 25 times greater, and signaling activity was directed toward all VQs. Although vibration signal activity was more pronounced in colonies with afterswarms, the signal was consistently associated with increased VQ survival under all conditions. These trends were exhibited similarly in the African and European colonies, suggesting that they have broad applicability to queen-replacement decisions over a range of environmental and racial conditions. However, the African and European colonies differed in the total number of queens involved in the elimination process and the relative importance of queen duels and pre-emergence destruction under the different reproductive strategies. Taken together, our results suggest that worker behavior is a major determinant for the outcome of queen replacement, either through reduced interactions that allow first-emerged queens to rapidly eliminate rivals, or through increased use of interactions such as the vibration signal, which may allow workers to influence the ultimate fate of each emerged VQ. We discuss the possibility that these behavior patterns may reflect the roles of cooperation and conflict in shaping honey bee reproductive decisions. Received 8 May 2007; revised 7 November 2007; accepted 20 November 2007.     

Influence of queen weight and colony origin on worker response in Solenopsis geminata

Abstract. The influence of weight and colony origin of the queen of Solenopsis geminata (F.) (Hymenoptera: Formicidae) on worker attraction is studied under laboratory conditions. In the first experiment, worker response to individual queens of different weight from the same colony is evaluated. Heavier queens are more attractive than smaller queens to their own workers. In subsequent experiments, the colony origin effect is investigated and worker response to a pair of queens of the same weight from the same or different colonies is compared. When queens are from the same colony, workers do not show a significant preference between queens. However, when queens are from a different colony, workers are significantly more attracted to their own queen than to the foreign queen. Finally, the response of workers to queens of different weight from the same or different colonies is investigated. In both cases, workers are significantly more attracted to a heavier queen than a lighter queen, even if the lighter queen is their own queen. A putative pheromonal component (E)‐6‐(1‐pentenyl)‐2H‐2‐pyranone, is not positively correlated with queen weight.     

Queen attendance behavior of worker honey bees (Apis mellifera L.) that are high and low responding to queen mandibular pheromone

Summary Queen attendance behavior of workers from selected honey bee colonies with high and low worker retinue response to synthetic queen mandibular gland pheromone (QMP) was investigated. Antennating, licking, grooming, and feeding of the queen by workers from high and low responding colonies were examined. High and low QMP responding workers did not attend the queen differently. However, workers originating from different colonies antennated and licked the queen more frequently than others, suggesting there may be a genetic basis for queen attendance behavior not necessarily associated with response to QMP. The median age of queen attendance was independent of strain.     

The role of heliothine hairpencil compounds in female Heliothis virescens (Lepidoptera: Noctuidae) behavior and mate acceptance

Studies on numerous insect species suggest that male-produced sex pheromones play a role in attracting females; as aphrodisiacs, making females more quiescent; or as a means of inhibiting competing males. Male heliothine moths display abdominal hairpencils during courtship, but the specific effects of the odors released on female behavior have not yet been elucidated. This study investigates the role of male hairpencil compounds in female Heliothis virescens mating behavior. Female H. virescens were exposed to filter paper loaded with hairpencil extracts of male H. virescens, Heliothis subflexa and Helicoverpa zea, and observed for behavioral responses to odors. Single synthetic compounds found in the H. virescens hairpencil blend were also tested. In mating assays between single male and female H. virescens it was found that: (i) antennectomized females mated less frequently than sham-operated females; (ii) females mated less frequently with males whose hairpencils had been surgically removed; (iii) females mated with males with ablated hairpencils if a filter paper loaded with one male equivalent of H. virescens hairpencil extract was presented simultaneously; and (iv) this effect was species-specific, as presentation of H. subflexa or H. zea hairpencil extracts did not restore mate acceptance. This study suggests that odors released by male hairpencils are important in mate acceptance by female H. virescens, and may play a role in mate choice and species isolation.     

蜜蜂蜂王信息素研究进展

综述了蜂王信息素的化学组成、特性、作用 ,以及与幼虫信息相互关系的研究进展 ,对蜂王信息素在养蜂和植物授粉中的应用做了介绍。西方蜜蜂的蜂王上颚腺信息素 (queen’smandibularglandpheromone,QMP)。QMP包含了 5种成分 :(E) -9-氧 -2 -癸烯酸 ( 9 ODA)、(R ,E) ( ) 和 (S,E) ( +) 9 羟基 -2 -癸烯酸 ( -9 HDA和 +9 HDA)、甲基p -羟基安息香酸盐 (HOB)、4-羟基 -3 -甲氧苯乙醇 (也称香草醇 ,HVA) ,9 ODA是其中最重要的成分 ;QMP的组分与蜜蜂的进化程度有关 ,进化程度越高则组分越复杂 ;QMP通过抑制保幼激素的产生来调节青年工蜂的个体发育。     

The Releaser Effects of the Tergal Gland Secretion of Queen Honeybees (Apis mellifera)

Queen honeybees are attractive to their workers, due partially to the pheromonal bouquet they secrete. Queen mandibular gland pheromone is a powerful attractant to worker honeybees but it is not solely responsible for eliciting retinue behavior. The attractiveness of virgin queen tergal gland secretions and queen mandibular pheromone to worker honeybees was tested using a retinue bioassay. The number of workers attending the treated pseudoqueen lures was videorecorded in order to allow for the quantification of attractiveness. Queen mandibular gland secretions were more attractive than tergal gland secretions (P 0.008), and both queen tergal gland secretions (P 0.0002) and mandibular gland secretions (P 0.0001) were significantly more attractive than the control treatment. This laboratory bioassay indicates that queen tergal gland secretions have a releaser effect that evokes retinue behavior from worker honeybees.     

Advertised quality,caste and food availability influence the survival cost of juvenile hormone in paper wasps

Life-history trade-offs are often hormonally mediated. Here, we provide a comparative perspective on the endocrine basis of life-history trade-offs by examining the invertebrate hormone juvenile hormone (JH). JH is often associated with benefits, including increased dominance and reproductive success. We tested whether JH reduced survival of Polistes dominulus wasps and whether this survival cost was influenced by factors such as advertised quality, food availability, caste and body size. Overall, JH reduced individual survival. Among fed and unfed queens, JH reduced survival in a dose-dependent manner. Among workers, JH had a stronger effect on survival of fed workers than unfed workers. Unfed workers died quickly and body size was the best predictor of survival. Surprisingly, queens and workers treated with JH survived longer when they had signals advertising high quality than when they had signals advertising low quality. The relationship between advertised quality and ability to withstand high levels of JH suggests that there are differential physiological costs associated with ornament elaboration that could play a role in maintaining signal accuracy over evolutionary time. Overall, the convergence of endocrine-mediated costs across diverse systems suggests that endocrine-mediated trade-offs may be an adaptive way to optimize resource allocation rather than a non-adaptive constraint specific to a particular hormone.