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Several animals, including bees, use visual search to distinguish targets of interest and ignore distractors. While bee flower choice is well studied, we know relatively little about how they choose between multiple rewarding flowers in complex floral environments. Two factors that could influence bee visual search for multiple flowers are the saliency (colour contrast against the background) and the reward value of flowers. We here investigated how these two different factors contribute to bee visual search. We trained bees to independently recognize two rewarding flower types that, in different experiments, differed in either saliency, reward value or both. We then measured their choices and attention to these flowers in the presence of distractors in a test without reinforcement. We found that bees preferred more salient or higher rewarding flowers and ignored distractors. When the high-reward flowers were less salient than the low-reward flowers, bees were nonetheless equally likely to choose high-reward flowers, for the reward and saliency values we used. Bees were also more likely to attend to these high-reward flowers, spending higher inspection times around them and exhibiting faster search times when choosing them. When flowers differed in reward, we also found an effect of the training order with low-reward targets being more likely to be chosen if they had been encountered during the more immediate training session prior to the test. Our results parallel recent findings from humans demonstrating that reward value can attract attention even when targets are less salient and irrelevant to the current task. [ABSTRACT FROM AUTHOR]

Flying safely and avoiding obstacles in low light is crucial for the bumblebees that forage around dawn and dusk. Previous work has shown that bumblebees overcome the limitations of their visual system—typically adapted for bright sunlight—by increasing the time over which they sample photons. While this improves visual sensitivity, it decreases their capacity to resolve fast motion. This study investigates what effect this has on obstacle avoidance in flight, a task that requires the bees to reliably detect obstacles in the frontal visual field and to make a timely diversion to their flight path. In both bright and dim light, bumblebees avoided the 5 cm diameter obstacle at a consistent distance (22 cm) although in dim light they approached it more slowly from a distance of at least at least 80 cm. This suggests that bumblebees have an effective strategy for avoiding obstacles in all light conditions under which they are naturally active, and it is hypothesised that this is based on a time-to-contact prediction. [ABSTRACT FROM AUTHOR]

This study examines the mechanism underlying one way in which bumblebees are known to develop a preference for symmetric patterns: through prior non-differential reinforcement on simple patterns (black discs and white discs). In three experiments, bees were given a choice among symmetric and asymmetric black-and-white non-rewarding patterns presented at the ends of corridors in a radial maze. Experimental groups had prior rewarded non-discrimination training on white patterns and black patterns, while control groups had no pre-test experience outside the colony. No preference for symmetry was obtained for any of the control groups. Prior training with circular patterns highlighting a horizontal axis of symmetry led to a specific subsequent preference for horizontal over vertical symmetry, while training with a vertical axis abolished this effect. Circles highlighting both axes created a general avoidance of asymmetry in favour of symmetric patterns with vertical, horizontal or both axes of symmetry. Training with plain circles, but not with deformed circles, led to a preference for symmetry: there was no evidence that the preference emerged just by virtue of having attention drawn away from irrelevant pattern differences. Our results point to a preference for symmetry developing gradually through first learning to extract an axis of symmetry from simple patterns and subsequently recognizing that axis in new patterns. They highlight the importance of continued learning through non-differential reinforcement by skilled foragers. Floral guides can function not only to guide pollinators to the source of reward but also to highlight an axis of symmetry for use in subsequent floral encounters. [ABSTRACT FROM AUTHOR]

We trained worker bumblebees to discriminate arrays of artificial nectaries (one, two, and three microcentrifuge tubes inserted into artificial flowers) from which they could forage in association with their location in a three-compartmental maze. Additionally, we challenged bees to learn to accomplish three different tasks in a fixed sequence during foraging. To enter the main three-compartmented foraging arena, they had first to slide open doors in an entry box to be able to proceed to an artificial flower patch in the main arena where they had to lift covers to the artificial nectaries from which they then fed. Then, the bees had to return to the entrance way to their hive, but to actually enter, were challenged to rotate a vertically oriented disc to expose the entry hole. The bees were adept at associating the array of nectaries with their position in the compartmental maze (one nectary in compartment one, two in two, and three in three), taking about six trials to arrive at almost error-free foraging. Over all it took the bees three days of shaping to become more or less error free at the multi-step suite of sequential task performances. Thus, they had learned where they were in the chain sequence, which array and in which compartment was rewarding, how to get to the rewarding array in the appropriate compartment, and finally how to return as directly as possible to their hive entrance, open the entrance, and re-enter the hive. Our experiments were not designed to determine the specific nature of the cues the bees used, but our results strongly suggest that the tested bees developed a sense of subgoals that needed to be achieved by recognizing the array of elements in a pattern and possibly chain learning in order to achieve the ultimate goal of successfully foraging and returning to their colony. Our results also indicate that the bees had organized their learning by a hierarchy as evidenced by their proceeding to completion of the ultimate goal without reversing their foraging paths so as to return to the colony without food. [ABSTRACT FROM AUTHOR]

During foraging, worker bumblebees are challenged by simple to complex tasks. Our goal was to determine whether bumblebees could successfully accomplish tasks that are more complex than those they would naturally encounter. Once the initial training to successfully manipulate a simple, artificial flower was completed, the bees were either challenged with a series of increasingly difficult tasks or with the most difficult task without the opportunity for prior learning. The first experiment demonstrated that the bees learned to slide or lift caps that prevented their access to the reinforcer sugar solution through a series of tasks with increasing complexity: moving one cap either to the right or to the left, or lifting it up. The second experiment demonstrated that the bees learned to push balls of escalating masses (diameters 1 and 1.27 cm) from the access to the hidden rewarding (sugar syrup) reservoir of artificial flowers. In both experiments, when bees with experience with only the simplest task (i.e. an artificial flower without a barrier to the reinforcer) were presented next with the most complex or difficult task, they failed. Only by proceeding through the series of increasingly difficult tasks were they able to succeed at the most difficult. We also noted idiosyncratic behaviours by individual bees in learning to succeed. Our results can be interpreted within the context of Skinnerian shaping and possibly scaffold learning. [ABSTRACT FROM AUTHOR]

Studies of bee cognition frequently use two-dimensional stimuli referred to as floral patterns, and yet how bees perceive pictorial representations is not known. An investigation of bumblebee ( Bombus impatiens) picture-object correspondence was undertaken according to the theory of Fagot et al. (Picture perception in animals. Psychology Press Ltd, East Sussex, pp 295-320, ) that pictures and objects may be confused, perceived as independent or equivalent. In three experiments, bumblebees were given discrimination training and unrewarded testing in a radial maze. In the first experiment, preferences between artificial flowers and photographs of those flowers revealed a significant learned preference for the stimulus rewarded during training: no confusion following training. In the second experiment, bees did transfer learning from artificial flowers to photographs: some commonality between an object and photograph was perceived. In the third experiment, bees spontaneously generalized a learned preference for one artificial flower to its silhouette, but only for one of two flowers used in training. No generalization was obtained to drawn images. Some transfer between image and object is exhibited, likely by low-level feature matching, but transfer is poor with degraded images, cautioning against extrapolation of picture-based responding to natural correspondents. Despite evidence that bees exhibit some transfer while retaining discrimination, it is likely that the observed response is due to generalization more akin to confusion than true equivalence. Furthermore, although 2D patterning cues (line, edge and shade) provide discriminable cues for bees between 2D stimuli, it is not here supported that such features are perceived as equivalent to the intended floral structures. [ABSTRACT FROM AUTHOR]

If animals are trained with two similar stimuli such that one is rewarding (S+) and one punishing (S−), then following training animals show a greatest preference not for the S+, but for a novel stimulus that is slightly more different from the S− than the S+ is. This peak shift phenomenon has been widely reported for vertebrates and has recently been demonstrated for bumblebees and honey bees. To explore the nature of peak shift in invertebrates further, here we examined the properties of peak shift in honey bees trained in a free-flight olfactory learning assay. Hexanal and heptanol were mixed in different ratios to create a continuum of odour stimuli. Bees were trained to artificial flowers such that one odour mixture was rewarded with 2 molar sucrose (S+), and one punished with distasteful quinine (S−). After training, bees were given a non-rewarded preference test with five different mixtures of hexanal and heptanol. Following training bees' maximal preference was for an odour mixture slightly more distinct from the S− than the trained S+. This effect was not seen if bees were initially trained with two distinct odours, replicating the classic features of peak shift reported for vertebrates. We propose a conceptual model of how peak shift might occur in honey bees. We argue that peak shift does not require any higher level of processing than the known olfactory learning circuitry of the bee brain and suggest that peak shift is a very general feature of discrimination learning. [ABSTRACT FROM AUTHOR]

Bumblebees are capable of rapidly learning discriminations, but flexibility in bumblebee learning is less well understood. We tested bumblebees ( Bombus impatiens) on a serial reversal learning task. A serial reversal task requires learning of an initial discrimination between two differentially rewarded stimuli, followed by multiple reversals of the reward contingency between stimuli. A reduction in errors with repeated reversals in a serial reversal task is an indicator of behavioural flexibility. Bees were housed in a large indoor environment and tested during foraging flights. Testing free-flying bees allowed for large numbers of trials and reversals. All bees were trained to perform a simultaneous discrimination between two colours for a nectar reward, followed by nine reversals of this discrimination. Results showed that bumblebees reduced errors and improved their performance across successive reversals. A reduction in perseverative errors was the major cause of the improvement in performance. Bees showed a slight increase in error rate in their final trials, perhaps as a consequence of increasing proactive interference, but proactive interference may also have contributed to the overall improvement in performance across reversals. Bumblebees are thus capable of behavioural flexibility comparable to that of other animals and may use proactive interference as a mechanism of behavioural flexibility in varying environments. [ABSTRACT FROM AUTHOR]

The capacity to discriminate between choice options is crucial for a decision-maker to avoid unprofitable options. The physical properties of rewards are presumed to be represented on context-dependent, nonlinear cognitive scales that may systematically influence reward expectation and thus choice behavior. In this study, we investigated the discrimination performance of free-flying bumblebee workers ( Bombus impatiens) in a choice between sucrose solutions with different concentrations. We conducted two-alternative free choice experiments on two B. impatiens colonies containing some electronically tagged bumblebees foraging at an array of computer-automated artificial flowers that recorded individual choices. We mimicked natural foraging conditions by allowing uncertainty in the probability of reward delivery while maintaining certainty in reward concentration. We used a Bayesian approach to fit psychometric functions, relating the strength of preference for the higher concentration option to the relative intensity of the presented stimuli. Psychometric analysis was performed on visitation data from individually marked bumblebees and pooled data from unmarked individuals. Bumblebees preferred the more concentrated sugar solutions at high stimulus intensities and showed no preference at low stimulus intensities. The obtained psychometric function is consistent with reward evaluation based on perceived concentration contrast between choices. We found no evidence that bumblebees reduce reward expectations upon experiencing non-rewarded visits. We compare psychometric function parameters between the bumblebee B. impatiens and the flower bat Glossophaga commissarisi and discuss the relevance of psychophysics for pollinator-exerted selection pressures on plants. [ABSTRACT FROM AUTHOR]

Bumblebees were exposed to a discrimination procedure in which reinforcement was contingent on choice of one of two spatial locations. The correct choice depended on whether a stimulus display contained two identical stimuli or two different stimuli. Some bees were trained with color stimuli and tested with line grating stimuli and others with the opposite arrangement. Four colonies of bumblebees produced more correct than incorrect choices to both identical and different stimuli during the transfer phase. This pattern of results is a signature of choices under control of an identity ('same/different') concept. The results therefore indicate the existence of an identity concept in bumblebees. [ABSTRACT FROM AUTHOR]

In honeybees, the conditioning of the proboscis extension response (PER) has provided a powerful tool to explore the mechanisms underlying olfactory learning and memory. Unfortunately, PER conditioning does not work well for visual stimuli in intact honeybees, and performance is improved only after antennal amputation, thus limiting the analysis of visual learning and multimodal integration. Here, we study visual learning using the PER protocol in harnessed bumblebees, which exhibit high levels of odor learning under restrained conditions. We trained bumblebees in a differential task in which two colors differed in their rewarding values. We recorded learning performance as well as response latency and accuracy. Bumblebees rapidly learned the task and discriminated the colors within the first two trials. However, performance varied between combinations of colors and was higher when blue or violet was associated with a high reward. Overall, accuracy and speed were negatively associated, but both components increased during acquisition. We conclude that PER conditioning is a good tool to study visual learning, using Bombus impatiens as a model, opening new possibilities to analyze the proximate mechanisms of visual learning and memory, as well as the process of multimodal integration and decision-making. [ABSTRACT FROM AUTHOR]

Bumblebees ( Bombus spp.) foraging in the field typically reject flowers where they detect the olfactory footprints of previous visitors and hence avoid recently emptied inflorescences. A growing number of studies have begun to illustrate that associative learning shapes the development of this process, in both bumblebees and other bee species. This raises the question of what the default response to such marks is, but little is known about how inexperienced foragers use social information. Here, we offered flower-naive bees a choice between scent-marked flowers and unmarked alternatives and found that individuals neither avoided nor preferred marked flowers. Our findings provide no support for 'hard-wired' responses to scent marks in bumblebees and highlight the importance of associative learning in shaping social information use to match local circumstances. [ABSTRACT FROM AUTHOR]

We presented bumblebees a spatial memory task similar to that used with other species (e.g., cats, dogs, and pigeons). In some conditions we allowed for presence of scent marks in addition to placing local and global spatial cues in conflict. Bumblebees ( Bombus impatiens) were presented an array of artificial flowers within a flight cage, one flower offering reward (S+), while the others were empty (S−). Bees were tested with empty flowers. In Experiment 1, flowers were either moved at the time of testing or not. Bees returned to the flower in the same absolute position of the S+ (the flower-array-independent (FAI) position), even if it was in the wrong position relative to the S− and even when new flower covers prevented the use of possible scent marks. New flower covers (i.e., without possible scent marks) had the effect of lowering the frequency of probing behavior. In Experiment 2, the colony was moved between training and testing. Again, bees chose the flower in the FAI position of the S+, and not the flower that would be chosen using strictly memory for a flight vector. Together, these experiments show that to locate the S+ bees did not rely on scent marks nor the positions of the S−, though the S− were prominent objects close to the goal. Also, bees selected environmental features to remember the position of the S+ instead of relying upon a purely egocentric point of view. Similarities with honeybees and vertebrates are discussed, as well as possible encoding mechanisms. [ABSTRACT FROM AUTHOR]