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.Modern views on associative learning acknowledge that classical as well as instrumental conditioning depend upon associations between external cues or behavioral responses and internal representations of reward (Rescorla 1987). Within this context, the term "expectation," or "expectancy," denotes an activation of an internal representation of reward in the absence of reinforcement by the cues and events predicting such a reward (Tolman 1959;Logan 1960). According to theory, the reward value associated with a stimulus is not a static, intrinsic property of the stimulus. Thus, for example, animals can assign different appetitive values to a stimulus as a function of both their internal state at the time when the stimulus is encountered and the background of their previous experience with such stimulus. This means that specific neural mechanisms have evolved not only to detect the presence of reward but also to predict its occurrence and magnitude based on internal representations from past experiences, in turn activated by the subject's current motivational status (Schultz 2000).Studying this form of learning is critical for understanding how reward controls behavior, how it leads to the formation of reward expectations, and how the brain uses reward-related information to control goal-directed behavior. Studies on reward expectations, however, sometimes appear to be paradoxical in assessing the cognitive complexity underlying such processes, as well as the basic principles of planning and decision-making. The reason is probably to be found in the fact that an anticipatory imagery or idea aroused by learned associations is thought to underlie these phenomena. In principle, however, neither highly complex cognitive abilities nor consciousness phenomena are assumed to be the base...