RESUMEN
Communication is a fundamental feature of animal societies and helps their members to solve the challenges they encounter, from exploiting food sources to fighting enemies or finding a new home. Eusocial bees inhabit a wide range of environments and they have evolved a multitude of communication signals that help them exploit resources in their environment efficiently. We highlight recent advances in our understanding of bee communication strategies and discuss how variation in social biology, such as colony size or nesting habits, and ecological conditions are important drivers of variation in communication strategies. Anthropogenic factors, such as habitat conversion, climate change, or the use of agrochemicals, are changing the world bees inhabit, and it is becoming clear that this affects communication both directly and indirectly, for example by affecting food source availability, social interactions among nestmates, and cognitive functions. Whether and how bees adapt their foraging and communication strategies to these changes represents a new frontier in bee behavioral and conservation research.
Asunto(s)
Aclimatación , Ecosistema , Animales , Abejas , ComunicaciónRESUMEN
Regulation of pollen and nectar foraging in honeybees is linked to differences in the sensitivity to the reward. Octopamine (OA) participates in the processing of reward-related information in the bee brain, being a candidate to mediate and modulate the division of labour among pollen and nectar foragers. Here we tested the hypothesis that OA affects the resource preferences of foragers. We first investigated whether oral administration of OA is involved in the transition from nectar to pollen foraging. We quantified the percentage of OA-treated bees that switched from a sucrose solution to a pollen feeder when the sugar concentration was decreased experimentally. We also evaluated if feeding the colonies sucrose solution containing OA increases the rate of bees collecting pollen. Finally, we quantified OA and tyramine (TYR) receptor genes expression of pollen and nectar foragers in different parts of the brain, as a putative mechanism that affects the decision-making process regarding the resource type collected. Adding OA in the food modified the probability that foragers switch from nectar to pollen collection. The proportion of pollen foragers also increased after feeding colonies with OA-containing food. Furthermore, the expression level of the AmoctαR1 was upregulated in foragers arriving at pollen sources compared with those arriving at sugar-water feeders. Using age-matched pollen and nectar foragers that returned to the hive, we detected an upregulated expression of a TYR receptor gene in the suboesophageal ganglia. These findings support our prediction that OA signalling affects the decision in honeybee foragers to collect pollen or nectar.
Asunto(s)
Conducta Animal/fisiología , Encéfalo/metabolismo , Conducta Alimentaria/fisiología , Expresión Génica/fisiología , Animales , Abejas , Néctar de las Plantas/metabolismo , Polen/metabolismo , Receptores de Amina Biogénica/metabolismo , Sacarosa/metabolismoRESUMEN
Plants and pollinators form beneficial relationships, with plants offering resources in return for pollination services. Some plants, however, add compounds to nectar to manipulate pollinators. Caffeine is a secondary plant metabolite found in some nectars that affects foraging in pollinators. In honeybees, caffeine increases foraging and recruitment to mediocre food sources, which might benefit the plant, but potentially harms the colonies. For the largest group of social bees, the stingless bees, the effect of caffeine on foraging behaviour has not been tested yet, despite their importance for tropical ecosystems. More generally, recruitment and foraging dynamics are not well understood in most species. We examined whether caffeine affects the foraging behaviour of the stingless bee Plebeia droryana, which frequently visits plants that produce caffeinated nectar and pollen. We trained bees to food sources containing field-realistic concentrations of sugar and caffeine. Caffeine did not cause P. droryana to increase foraging frequency and persistence. We observed P. droryana recruiting to food sources; however, this behaviour was also not affected by caffeine. Instead we found that higher sugar concentrations caused bees to increase foraging effort. Thus, unlike in other pollinators, foraging behaviour in this stingless bee is not affected by caffeine. As the Brazilian P. droryana population that we tested has been exposed to coffee over evolutionary time periods, our results raise the possibility that it may have evolved a tolerance towards this central nervous system stimulant. Alternatively, stingless bees may show physiological responses to caffeine that differ from those of other bee groups.
Asunto(s)
Abejas/fisiología , Variación Biológica Individual , Cafeína/metabolismo , Azúcares/metabolismo , Animales , Conducta Apetitiva , Brasil , Conducta Alimentaria , Conducta SocialRESUMEN
Many ant and termite colonies are defended by soldiers with powerful mandibles or chemical weaponry. Recently, it was reported that several stingless bee species also have soldiers for colony defence. These soldiers are larger than foragers, but otherwise lack obvious morphological adaptations for defence. Thus, how these soldiers improve colony fitness is not well understood. Robbing is common in stingless bees and we hypothesized that increased body size improves the ability to recognize intruders based on chemosensory cues. We studied the Neotropical species Tetragonisca angustula and found that large soldiers were better than small soldiers at recognizing potential intruders. Larger soldiers also had more olfactory pore plates on their antennae, which is likely to increase their chemosensory sensitivity. Our results suggest that improved enemy recognition might select for increased guard size in stingless bees.
Asunto(s)
Abejas/fisiología , Comportamiento de Nidificación , Sensilos/anatomía & histología , Agresión , Animales , Abejas/anatomía & histología , Tamaño Corporal , Olfato , Conducta SocialRESUMEN
The differentiation of workers into morphological castes represents an important evolutionary innovation that is thought to improve division of labor in insect societies. Given the potential benefits of task-related worker differentiation, it is puzzling that physical worker castes, such as soldiers, are extremely rare in social bees and absent in wasps. Following the recent discovery of soldiers in a stingless bee, we studied the occurrence of worker differentiation in 28 stingless bee species from Brazil and found that several species have specialized soldiers for colony defence. Our results reveal that worker differentiation evolved repeatedly during the last ~ 25 million years and coincided with the emergence of parasitic robber bees, a major threat to many stingless bee species. Furthermore, our data suggest that these robbers are a driving force behind the evolution of worker differentiation as targets of robber bees are four times more likely to have nest guards of increased size than non-targets. These findings reveal unexpected diversity in the social organization of stingless bees.Although common in ants and termites, worker differentiation into physical castes is rare in social bees and unknown in wasps. Here, Grüter and colleagues find a guard caste in ten species of stingless bees and show that the evolution of the guard caste is associated with parasitization by robber bees.
Asunto(s)
Abejas , Conducta Animal , Evolución Biológica , Animales , Femenino , Masculino , Agresión/fisiología , Abejas/anatomía & histología , Abejas/clasificación , Abejas/fisiología , Conducta Animal/fisiología , Tamaño Corporal , Brasil , FilogeniaRESUMEN
Division of labor among workers is common in insect societies and is thought to be important in their ecological success. In most species, division of labor is based on age (temporal castes), but workers in some ants and termites show morphological specialization for particular tasks (physical castes). Large-headed soldier ants and termites are well-known examples of this specialization. However, until now there has been no equivalent example of physical worker subcastes in social bees or wasps. Here we provide evidence for a physical soldier subcaste in a bee. In the neotropical stingless bee Tetragonisca angustula, nest defense is performed by two groups of guards, one hovering near the nest entrance and the other standing on the wax entrance tube. We show that both types of guards are 30% heavier than foragers and of different shape; foragers have relatively larger heads, whereas guards have larger legs. Low variation within each subcaste results in negligible size overlap between guards and foragers, further indicating that they are distinct physical castes. In addition, workers that remove garbage from the nest are of intermediate size, suggesting that they might represent another unrecognized caste. Guards or soldiers are reared in low but sufficient numbers (1-2% of emerging workers), considering that <1% usually perform this task. When challenged by the obligate robber bee Lestrimelitta limao, an important natural enemy, larger workers were able to fight for longer before being defeated by the much larger robber. This discovery opens up opportunities for the comparative study of physical castes in social insects, including the question of why soldiers appear to be so much rarer in bees than in ants or termites.
Asunto(s)
Adaptación Biológica/fisiología , Abejas/anatomía & histología , Abejas/fisiología , Jerarquia Social , Comportamiento de Nidificación/fisiología , Conducta Social , Conducta Agonística/fisiología , Animales , Pesos y Medidas Corporales , Brasil , Modelos Lineales , ObservaciónRESUMEN
The honeybee (Apis mellifera) waggle dance is one of the most intriguing animal communication signals. A dancing bee communicates the location of a profitable food source and its odour. Followers may often experience situations in which dancers indicate an unfamiliar location but carry the scent of a flower species the followers experienced previously at different locations. Food scents often reactivate bees to resume food collection at previously visited food patches. This double function of the dance creates a conflict between the social vector information and the private navigational information. We investigated which kind of information followers with field experience use in this situation and found that followers usually ignored the spatial information encoded by the waggle dance even if they followed a dance thoroughly (five waggle runs or more). They relied on private information about food source locations instead (in 93% of all cases). Furthermore, foragers preferred to follow dancers carrying food odours they knew from previous field trips, independently of the spatial information encoded in the dance. Surprisingly, neither odour identity nor the location indicated by the dancer was an important factor for the reactivation success of a dance. Our results contrast with the assumption that (i) followers usually try to decode the vector information and (ii) dances indicating an unfamiliar location are of little interest to experienced foragers.
Asunto(s)
Comunicación Animal , Abejas/fisiología , Animales , Conducta Alimentaria/fisiología , Alimentos , Odorantes , Conducta SocialRESUMEN
Recent studies showed that nectar odors brought back by honeybee foragers can be learned associatively inside the hive. In the present study, we focused on the learning abilities of bees, which directly interact via trophallaxis with the incoming nectar foragers: the workers that perform nectar-receiving tasks inside the hive. Workers that have received food directly from foragers coming back from a feeder offering either unscented or scented sugar solution [phenylacetaldehyde (PHE) or nonanal diluted] were captured from two observational hives, and their olfactory memories were tested using the proboscis extension response paradigm. Bees that have received scented solution from incoming foragers showed significantly increased response frequencies for the corresponding solution odor in comparison with those that have received unscented solution. No differences in the response frequencies were found between food odors and colonies. The results indicate that first-order receivers learn via trophallaxis the association between the scent and the sugar solution transferred by incoming foragers. The implications of these results should be considered at three levels: the operational cohesion of bees involved in foraging-related tasks, the information propagation inside the hive related to the floral type exploited, and the putative effect of these memories on future preferences for resources.
Asunto(s)
Abejas/fisiología , Conducta Alimentaria , Flores , Odorantes , Olfato/fisiología , Alimentación Animal , Animales , Conducta SocialRESUMEN
A honeybee hive serves as an information centre in which communication among bees allows the colony to exploit the most profitable resources in a continuously changing environment. The best-studied communication behaviour in this context is the waggle dance performed by returning foragers, which encodes information about the distance and direction to the food source. It has been suggested that another information cue, floral scents transferred within the hive, is also important for recruitment to food sources, as bee recruits are more strongly attracted to odours previously brought back by foragers in both honeybees and bumble-bees. These observations suggested that honeybees learn the odour from successful foragers before leaving the hive. However, this has never been shown directly and the mechanisms and properties of the learning process remain obscure. We tested the learning and memory of recruited bees in the laboratory using the proboscis extension response (PER) paradigm, and show that recruits indeed learn the nectar odours brought back by foragers by associative learning and retrieve this memory in the PER paradigm. The associative nature of this learning reveals that information was gained during mouth-to-mouth contacts among bees (trophallaxis). Results further suggest that the information is transferred to long-term memory. Associative learning of food odours in a social context may help recruits to find a particular food source faster.