Serial reversal learning in bumblebees (Bombus impatiens)

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.     

Decision-making and associative color learning in harnessed bumblebees (Bombus impatiens)

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.     

Spatial encoding by bumblebees (Bombus impatiens) of a reward within an artificial flower array

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.     

Transposition of flower height by bumble bee foragers (Bombus impatiens)

Honeybees learn visual characteristics of reinforced and non-reinforced flowers in differential conditioning experiments (i.e., experiments that require subjects to choose between a reinforced and non-reinforced flower). In this study bumble bee foragers (Bombus impatiens) were trained in a transposition paradigm to determine if the relational properties of flowers also influence choice behavior. Subjects in one group (group A) were trained with repeated choices between a medium-height flower replete with sucrose solution and an empty tall flower until the medium-height flower was sampled preferentially in five consecutive trials. A second group (group B) was trained on the medium height flower alone for five trials. In a single test trial subjects were given a choice between the medium-height flower and a short flower, each filled with water. A control group showed no preference in this test. Group B subjects showed a significant preference for the medium-height flower and group A subjects that exhibited flower constancy (i.e., sampled only the medium-height flower in training trials) showed an identical pattern of choice. Subjects in group A that sampled both flowers during training, in contrast, transposed flower height and preferred the short flower. These results suggest that the choice behavior of bumble bee foragers is influenced by relational and absolute properties of flowers. The flower characteristics learned by foragers appear to depend on the difficulty of the discrimination problem and the context in which flowers are sampled. Patches of flowers limited in phenotypic variability may produce simple associative learning and flower constancy by foragers. Received: 17 September 1999 / Accepted after revision: 22 June 2000     

Problem solving by men and mammals

There are two opposed theories which attempt to account for the processes of problem solution involved in learning and intelligence. The former is neural in its basis and postulates the existence of a bare connection as a bonding or linkage of two experiences. The second theory, that of gestalt, implies that learning or apprehension involves a relationship of the parts of the experience to each other as well as to the whole. While these psychological schools are exclusive of and opposed to each other, yet they are merely extremes of what actually exists. There is a minimum level of learning in which associationism is operative and a maximum intellectual level at which explicit relationships predominate. An examination of experimental results will show this to be the case. The results of conditioned reflex experiments in both animals and man appear to show that there is present nothing beyond arbitrary linkage or bonding between the parts of the situation involved. On the other hand, one finds a certain degree of implicit meaning involved in perceptual situations—the level of animal achievement—as well as a certain measure of transfer of meaning. Explicit meanings may be related by means of word symbols involving the principle of the concept through the medium of speech, and this is only arrived at by human beings. We may regard these higher levels as “emergents” from the level of bonding or linkage.     

Problem solving skills in young yellow-crowned parakeets (Cyanoramphus auriceps)

Despite the long divergent evolutionary history of birds and mammals, early avian and primate cognitive development have many convergent features. Some of these features were investigated with a series of tasks designed to assess human infant development. The tasks were presented to young parakeets to assess their means-end problem solving abilities. Examples of these early skills are: attaining and playing with objects, retrieving rewards through use of a stick or rake, or by pulling in rewards on supports or on the ends of strings. Twelve such tasks were presented to 11 young yellow-crowned parakeets (Cyanoramphus auriceps) to investigate their natural abilities; there was no attempt to train them to do those tasks that they did not spontaneously perform. Six of the birds were parent-raised and five were hand-raised. The birds completed 9 of the 12 tasks, demonstrating all the Piagetian sensorimotor circular reactions, but they failed to hand-watch ("claw-watch"), to stack objects, or to fill a container. Their ordinality on the tasks differed from that of human infants in that locomotion to obtain objects occurred earlier in the avian sequence of development and the mid-level tasks were performed by the two groups of avian subjects in a mixed order perhaps indicating that these abilities may not emerge in any particular order for these birds as they supposedly do for human infants. The hand-raised group needed fewer sessions to complete these means-end tasks. Electronic Publication     

Problem solving ability of Octopus vulgaris Lamarck (Mollusca, Cephalopoda)

Experiments presented in this study show that Octopus vulgaris Lamarck is able to open transparent glass jars closed with a plastic plug and containing a live crab (Carcinus mediterraneus). The animals remove the plus (Operandum: O) and seize the crab (Predation :P) in one single attack. The number of unsuccessful attacks appears to decrease over a series of trials (p less than .01); during the same period exploration time remains unchanged. There is a statistically significant increase in performance over trials for O (p less than .01) and P (p less than .05) mean times analyzed by single factor ANOVA, suggesting that the learning process is accomplished either by stimulus-response association or by trial and error. We propose that Octopus vulgaris is capable of learning the solutions of both problems, Operandum and Predation, thus showing a highly developed ability of "integration" of the behavioral program.     

Problem solving: Increased planning with practice

Two experiments using two isomorphs of the Tower of Hanoi show that participants increase the amount of planning they do as they learn that it increases problem solving efficiency. In addition, competition among different approaches emerged as participants gained more experience with the task, with an optimal strategy gradually replacing a less effective, though easier one. It is hypothesized that the competition among approaches is mediated by the costs incurred in terms of the number of moves needed to solve the problems. An ACT-R model of participant performance is used to validate this hypothesized mechanism and to examine many of the details of participants’ performance. This model corresponds closely to the observed data, from overall performance in terms of number of moves to details of participants’ strategy choices and variability in strategy use among individuals.     

Growth and pruning of mushroom body Kenyon cell dendrites during worker behavioral development in the paper wasp, Polybia aequatorialis (Hymenoptera: Vespidae)

Adult workers of some social insect species show dramatic behavioral changes as they pass through a sequence of task specializations. In the paper wasp, Polybia aequatorialis, female workers begin adult life within the nest tending brood, progress to maintaining and defending the nest exterior, and ultimately leave the nest to forage. The mushroom body (MB) calyx neuropil increases in volume as workers progress from in-nest to foraging tasks. In other social Hymenoptera (bees and ants), MB Kenyon cell dendrites, axons and synapses change with the transition to foraging, but these neuronal effects had not been studied in wasps. Furthermore, the on-nest worker of Polybia wasps, an intermediate task specialization not identified in bees or ants, provides the opportunity to study pre-foraging worker class transitions. We asked whether Kenyon cell dendritic arborization varies with the task specialization of Polybia workers observed in the field near Monteverde, Costa Rica. Golgi-impregnated arbors in the lip and collar calyces, which receive a predominance of olfactory and visual input, respectively, were quantified using Sholl’s concentric circles and a novel application of virtual spherical probes. Arbors of the lip varied in a manner reminiscent of honeybees, with foragers having the largest and in-nest workers having the smallest arbors. In contrast, arbors of the collar were largest in foragers but smallest in on-nest workers. Thus, progression through task specializations in P. aequatorialis involves subregion specific dendritic growth and regression in the MB neuropil. These results may reflect the sensitivity of Kenyon cell dendritic structure to specialization dependent social and sensory experience.     

Dissociable stages of problem solving (I): temporal characteristics revealed by eye-movement analyses

Understanding the functional neuroanatomy of planning and problem solving may substantially benefit from better insight into the chronology of the cognitive processes involved. Based on the assumption that regularities in cognitive processing are reflected in overtly observable eye-movement patterns, here we recorded eye movements while participants worked on Tower of London (TOL) problems that comprised an experimental manipulation of different task demands. Single-trial saccade-locked analyses revealed that higher demands on forming an internal problem representation were associated with an increased number of gaze alternations between start state and goal state, but did not show any effect on the durations of these inspections of the states. In contrast, higher demands on actual planning in terms of mental manipulations of working memory contents coincided with a prolonged duration of the very last inspection of the start state (i.e., immediately preceding movement execution) but did not show any effect on the number of gaze alterations. Differential task demands on internalization and planning processes during problem solving hence selectively affect different eye-movement parameters. Moreover, these findings complement previous neuroimaging data on dissociable contributions of left and right dorsolateral prefrontal cortex in problem solving with novel evidence for a corresponding dissociation in the eye-movement patterns reflecting the associated cognitive processes.     

Problem solving in semantically rich domains: An example from engineering thermodynamics

Recent research on human problem solving has largely focused on laboratory tasks that do not demand from the subject much prior, task-related information. This study seeks to extend the theory of human problem solving to semantically richer domains that are characteristic of professional problem solving. We discuss the behavior of a single subject solving problems in chemical engineering thermodynamics. We use as a protocol-encoding device a computer program called SAPA which also doubles as a theory of the subject's problem-solving behavior. The subject made extensive use of means-ends analysis, similar to that observed in semantically less rich domains, supplemented by recognition mechanisms for accessing information in semantic memory.     

Problem solving in infancy: the emergence of an action plan.

Young children's strategies were evaluated as they grasped and used objects. Spoons containing food and toys mounted on handles were presented to 9-, 14-, and 19-month-old children with the handle alternately oriented to the left and right. The alternating orientations revealed strategies that the children used for grasping items. Younger children usually reached with their preferred hand, disregarding the item's orientation. In the case of the spoon, this strategy produced awkward grasps that had to be corrected later. Older children anticipated the problem, alternated the hand used, and achieved an efficient radial grip (i.e., handle grasped with base of thumb toward food or toy end) for both orientations. A model of the development of action-selection strategies is proposed to illustrate planning in children younger than 2 years.