Learning reward expectations in honeybees.

The aim of this study was to test whether honeybees develop reward expectations. In our experiment, bees first learned to associate colors with a sugar reward in a setting closely resembling a natural foraging situation. We then evaluated whether and how the sequence of the animals' experiences with different reward magnitudes changed their later behavior in the absence of reinforcement and within an otherwise similar context. We found that the bees that had experienced increasing reward magnitudes during training assigned more time to flower inspection 24 and 48 h after training. Our design and behavioral measurements allowed us to uncouple the signal learning and the nutritional aspects of foraging from the effects of subjective reward values. We thus found that the animals behaved differently neither because they had more strongly associated the related predicting signals nor because they were fed more or faster. Our results document for the first time that honeybees develop long-term expectations of reward; these expectations can guide their foraging behavior after a relatively long pause and in the absence of reinforcement, and further experiments will aim toward an elucidation of the neural mechanisms involved in this form of learning.