Descripción: Early reports indicate that trophallaxis, i.e. the exchange of liquid food by mouth, may allow honeybees to assign nectar odours with predictive values to anticipate biological meaningful reward stimuli. Nevertheless, this type of learning has not been addressed directly. In the present study, pairs of animals were isolated to induce trophallaxis under controlled conditions and, afterwards, the honeybee proboscis extension reflex was used to investigate the possible role of trophallaxis in learning olfactory cues. The results demonstrate unambiguously that associative learning actually occurs by means of trophallaxis. Animals associate the odour (as the conditioned stimulus or CS) and the sucrose (as the unconditioned stimulus or US) present in the solution they receive through trophallaxis. Moreover, this particular kind of learning leads to long-term olfactory memories after a single learning trial, even when trophallaxis is brief. In addition, we found that the strength of association is clearly affected by CS and US intensity as well as the recent previous foraging experiences of the animals. Comparisons are presented among several features of the learning during trophallaxis and the classical conditioning of the proboscis extension reflex with restrained subjects. Finally, the relevance of learning through trophallaxis in the task of successful foraging is discussed.

Descripción: Camponotus mus ants can associate sucrose and odour at the source during successive foraging cycles and use this memory to locate the nectar in the absence of other cues. These ants perform conspicuous trophallactic behaviour during recruitment while foraging for nectar. In this work, we studied whether Camponotus mus ants are able to establish this odour-sucrose association in the social context of trophallaxis and we evaluated this memory in another context previously experienced by the ant, as a nectar source. After a single trophallaxis of a scented solution, the receiver ant was tested in a Y-maze without any reward, where two scents were presented: in one arm, the solution scent and in the other, a new scent. Ants consistently chose the arm with the solution scent and stayed longer therein. Trophallaxis duration had no effect on the arm choice or with the time spent in each arm. Workers are able to associate an odour (conditioned stimulus) with the sucrose (unconditioned stimulus) they receive through a social interaction and use this memory as choice criteria during food searching.

Descripción: Apis mellifera bees perform dances to communicate the presence of desirable nectar sources. The regulation of these dances does not depend exclusively on properties of the nectar sources, but also upon certain stimuli derived from the foraging status of the colony as a whole; i.e. bees exploiting a source of constant profitability are more likely to dance when the colony's nectar intake rate is low. Based on these stimuli, individual bees tune their dances according to their colony's nectar influx without visiting alternative nectar sources. Division of labour, in addition, is a common feature in honeybees. Upon returning to the nest, successful foragers transfer the content of their crops to food-receivers by means of a common behaviour in social insects called trophallaxis, i.e. the transfer of liquid food by mouth. Martin Lindauer stated that a returned forager may sense the foraging status of its colony on the basis of the food transfer process by computing how quickly and eagerly the food-receivers unload its crop. This study focuses on the forager's experience during the food transfer process, its variability based on the colony's nectar influx, and the separate effects that the 'ease' and the 'eagerness' of the food-unloading have on the tuning of recruitment dances. Results indicate that foragers can rapidly sense variations in the colony's nectar influx, even when they experience no variation in the time interval between their return to the hive and the beginning of the food transfer. To accomplish this task they appear to use stimuli derived from the number of food-receivers, which enable them, in turn, to set their dance thresholds in relation to the nectar influx of their colony. The relevance of these findings is discussed in the context of communication and successful foraging.

Descripción: Background: Honeybees (Apis mellifera) exhibit an extraordinarily tuned division of labor that depends on age polyethism. This adjustment is generally associated with the fact that individuals of different ages display different response thresholds to given stimuli, which determine specific behaviors. For instance, the sucrose-response threshold (SRT) which largely depends on genetic factors may also be affected by the nectar sugar content. However, it remains unknown whether SRTs in workers of different ages and tasks can differ depending on gustatory and olfactory experiences. Methodology: Groups of worker bees reared either in an artificial environment or else in a queen-right colony, were exposed to different reward conditions at different adult ages. Gustatory response scores (GRSs) and odor-memory retrieval were measured in bees that were previously exposed to changes in food characteristics. Principal Findings: Results show that the gustatory responses of pre-foraging-aged bees are affected by changes in sucrose solution concentration and also to the presence of an odor provided it is presented as scented sucrose solution. In contrast no differences in worker responses were observed when presented with odor only in the rearing environment. Fast modulation of GRSs was observed in older bees (12-16 days of age) which are commonly involved in food processing tasks within the hive, while slower modulation times were observed in younger bees (commonly nurse bees, 6-9 days of age). This suggests that older food-processing bees have a higher plasticity when responding to fluctuations in resource information than younger hive bees. Adjustments in the number of trophallaxis events were also found when scented food circulated inside the nest, and this was positively correlated with the differences in timing observed in gustatory responsiveness and memory retention for hive bees of different age classes. Conclusions: This work demonstrates the accessibility of chemosensory information in the honeybee colonies with respect to incoming nectar. The modulation of the sensory-response systems within the hive can have important effects on the dynamics of food transfer and information propagation. © 2010 Ramírez et al.

Descripción: Cognitive experiences during the early stages of life play an important role in shaping future behavior. Behavioral and neural long-term changes after early sensory and associative experiences have been recently reported in the honeybee. This invertebrate is an excellent model for assessing the role of precocious experiences on later behavior due to its extraordinarily tuned division of labor based on age polyethism. These studies are mainly focused on the role and importance of experiences occurred during the first days of the adult lifespan, their impact on foraging decisions, and their contribution to coordinate food gathering. Odor-rewarded experiences during the first days of honeybee adulthood alter the responsiveness to sucrose, making young hive bees more sensitive to assess gustatory features about the nectar brought back to the hive and affecting the dynamic of the food transfers and the propagation of food-related information within the colony. Early olfactory experiences lead to stable and long-term associative memories that can be successfully recalled after many days, even at foraging ages. Also they improve memorizing of new associative learning events later in life. The establishment of early memories promotes stable reorganization of the olfactory circuits inducing structural and functional changes in the antennal lobe (AL). Early rewarded experiences have relevant consequences at the social level too, biasing dance and trophallaxis partner choice and affecting recruitment. Here, we revised recent results in bees' physiology, behavior, and sociobiology to depict how the early experiences affect their cognition abilities and neural-related circuits. © 2013 Arenas, Ramírez, Balbuena and Farina.