Imitation and affordance learning by pigeons (Columba livia)

The bidirectional control procedure was used to determine whether pigeons (Columba livia) would imitate a demonstrator that pushed a sliding screen for food. One group of observers saw a trained demonstrator push a sliding screen door with its beak (imitation group), whereas 2 other groups watched the screen move independently (possibly learning how the environment works) with a conspecific either present (affordance learning with social facilitation) or absent (affordance learning alone). A 4th group could not see the screen being pushed (sound and odor control). Imitation was evidenced by the finding that pigeons that saw a demonstrator push the screen made a higher proportion of matching screen pushes than observers in 2 appropriate control conditions. Further, observers that watched a screen move without a demonstrator present made a significantly higher proportion of matching screen pushes than would be expected by chance. Thus, these pigeons were capable of affordance learning.     

Representation of serial order in pigeons (Columba livia)

In Experiment 1, 2 groups of pigeons were trained to respond to either a 4-item (A→B→C→D) or 5-item (A→B→C→D→E) list. After learning their respective list, half of the subjects were trained on a positive pair with reinforcement provided when pairs were responded to in the order true to that of the original sequence (4-item: B→C; 5-item: B→D). The remaining subjects were trained on a negative pair with reinforcement provided for responding to the pairs in the order opposite to that learned in the original sequence (4-item: C→B; 5-item: D→B). Subjects in the positive pair condition learned their respective pair faster than did subjects in the negative pair condition. In Experiment 2, after reaching criterion on a 4-item list, subjects received 16 BC probe trials spread across 4 sessions of training. Subjects performed significantly above chance on the probe trials. The performance of our subjects in Experiments 1 and 2 demonstrates that, similar to monkeys, pigeons form a representation of the lists that they learn.     

Imitation of conditional discriminations in pigeons (Columba livia)

In the present experiments, the 2-action method was used to determine whether pigeons could learn to imitate a conditional discrimination. Demonstrator pigeons (Columba livia) stepped on a treadle in the presence of 1 light and pecked at the treadle in the presence of another light. Demonstration did not seem to affect acquisition of the conditional discrimination (Experiment 1) but did facilitate its reversal of the conditional discrimination (Experiments 2 and 3). The results suggest that pigeons are not only able to learn a specific behavior by observing another pigeon, but they can also learn under which circumstances to perform that behavior. The results have implications for proposed mechanisms of imitation in animals.     

Flexible serial response learning by pigeons (Columba livia) and humans (Homo sapiens)

Experimental tasks designed to involve procedural memory are often rigid and unchanging, despite many reasons to expect that implicit learning processes can be flexible and support considerable variability. A version of the serial response time (SRT) task was developed, in which the locations of targets were probabilistically determined. Targets appeared in locations according to both a structured sequence and a cue validity parameter, and the time to respond to each target was measured. Pigeons (Columba livia) and humans (Homo sapiens) both showed response time facilitation at the highest tested value for cue validity, and the magnitude of that facilitation gradually weakened as cue validity was decreased. Both species showed evidence that response times were largely determined by the local predictabilities of individual cue locations. In addition, humans showed some evidence that explicit knowledge of the sequence affected response times, specifically when cue validity was 100%.     

Acquisition of a same-different discrimination task by pigeons (Columba livia)

Four homing pigeons were trained to discriminate two figures simultaneously presented on an LCD screen. The figure was either a rectangle (A) or a square (B), and four combinations of the two figures, AA, AB, BA, BB, appeared in a pseudo-randomized order. The pigeons' task was to peck one of these figures based upon whether the two figures were identical or not. One pigeon successfully learned this discrimination, with proportions of correct responses above 90% in two consecutive sessions. Of the other birds, two performed above chance level but had difficulty meeting a learning criterion of above 80% in two consecutive sessions. All birds achieved this criterion when the combinations of figures presented were reduced to two. Results suggested that learning the present same-different discrimination is within the capacity of pigeons to a certain extent, although there exists considerable individual variation in the pigeons' skills to acquire complex discrimination.     

Information processing by pigeons (Columba livia): incentive as information

Experiment 1 showed that the Hick-Hyman law (W. E. Hick, 1952; R. Hyman, 1953) described the effects of anticipated reinforcement, a form of incentive, on pigeons' (Columba livia) reaction time to respond to a target spatial location. Reaction time was an approximately linear function of amount of information interpreted as probability of reinforcement, implying that pigeons processed incentive at a constant rate. Experiment 2 showed that the Hick-Hyman law described effects of incentive even when it varied from moment to moment in a serial reaction time task similar to that of M. J. Nissen and P. Bullemer (1987), and processing information about target spatial location modulated absolute reaction time and not rate of processing incentive. The results support mental continuity and provide comparative support for the idea of the economics of information in economic theory about the incentive value of information.     

Multiple-landmark piloting in pigeons (Columba livia): landmark configuration as a discriminative cue

Using the landmark-transformation technique, researchers have shown that pigeons (Columba livia) tend to encode a goal location relative to 1 landmark, even when multiple landmarks are in the vicinity of the goal. The current experiments examined pigeons' ability to use configural information from a set of landmarks by making the arrangement of 4 landmarks a discriminative cue to the location of buried seeds. Results showed that pigeons used information from the 3 consistently placed landmarks to search accurately when 1 landmark was displaced. Findings indicate that pigeons are able to search for a goal using information from multiple landmarks instead of just 1 and that landmark use by these birds may be more flexible than previously theorized.     

Transfer from “edible” categorization training to feeding behavior in pigeons (Columba livia)1

Abstract: We investigated a transfer from an operant experimental situation to a feeding situation in pigeons using real objects as stimuli. Four pigeons were trained in an operant box to categorize familiar edible items as positives and inedible items as negatives with a go/no‐go procedure. Next, two pairs of unfamiliar edible items were added as stimuli. One of the paired stimuli was arbitrarily assigned as positive and the other as negative. We tested the subjects in their home cages to see whether they would feed on the items they were trained to categorize as positives. In three of the six cases in which categorization training was successful, they continued to peck the positive items. This result suggests that the pigeons transfer what they learned in the operant training situation to the feeding situation.     

Functional lateralization, interhemispheric transfer and position bias in serial reversal learning in pigeons (Columba livia)

In the present study we investigated lateralization of color reversal learning in pigeons. After monocular acquisition of a simple color discrimination with either the left or right eye, birds were tested in a serial reversal procedure. While there was only a slight and non-significant difference in choice accuracy during original color discrimination, a stable superiority of birds using the right eye emerged in serial reversals. Both groups showed a characteristic ‘learning-to-learn’ effect, but right-eyed subjects improved faster and reached a lower asymptotic error rate. Subsequent testing for interocular transfer demonstrated a difference between pre- and post-shift choice accuracy in pigeons switching from right to left eye but not vice versa. This can be accounted for by differences in maximum performance using either the left or right eye along with an equally efficient but incomplete interocular transfer in both directions. Detailed analysis of the birds’ response patterns during serial reversals revealed a preference for the right of two response keys in both groups. This bias was most pronounced at the beginning of a session. It decreased within sessions, but became more pronounced in late reversals, thus indicating a successful strategy for mastering the serial reversal task. Interocular transfer of response patterns revealed an unexpected asymmetry. Birds switching from right to left eye continued to prefer the right side, whereas pigeons shifting from left to right eye were now biased towards the left side. The results suggest that lateralized performance during reversal learning in pigeons rests on a complex interplay of learning about individual stimuli, stimulus dimensions, and lateralized response strategies. Received: 4 June 1999 / Accepted after revision: 18 August 1999     

Choice biases in no-sample and delay testing in pigeons (Columba livia)

Animal Cognition - In experimental tasks that involve stimuli that vary along a quantitative continuum, some choice biases are commonly found. Take, for instance, a matching-to-sample task where...     

Flexible learning and use of multiple-landmark information by pigeons (Columba livia) in a touch screen-based goal searching task

Three touch screen-based experiments were conducted to investigate whether pigeons would learn to use configural information about a goal's location in relation to a multiple-landmark array. In Experiment 1, 4 pigeons (Columba livia) were trained to peck a computer monitor at a location that constituted the third vertex of a hypothetical triangle defined by 2 different landmarks. The landmarks appeared in 3 orientations during the training, and the pigeons' goal-searching ability easily transferred to the landmarks presented in 3 novel orientations. Each landmark was asymmetric, so we next examined whether the pigeons used (a) the small-scale, local orientation information that could be inferred from each landmark individually, or (b) the large-scale, configural information that could be inferred from the spatial arrangement of multiple landmarks taken as a whole. Even when each single landmark appeared by itself, the pigeons were able to locate the goal accurately, suggesting that the large-scale, configural information was not essential. However, when 1 landmark locally pointed to a location that was consistent with the triangular configuration and the other landmark locally pointed to a different location, the pigeons predominantly pecked at the configurally array-consistent location. These results suggest that the pigeons redundantly learned both the large-scale, configural strategy and the local, single-landmark strategy, but they mainly used the latter information, and used the former information solely to disambiguate conflicts when the 2 landmarks pointed toward different targets. Such flexible learning and use of redundant information may reflect the pigeons' adaptation to unstable wild environments during their evolutionary history.     

Degree of representation of the matching concept in pigeons (Columba livia)

In Experiment 1, 12 pigeons (Columba livia) were trained on a simultaneous matching-to-sample task with 2 stimuli and then tested with 2 novel stimuli. Half of the birds were trained with a fixed ratio schedule requirement of 1 (FR1) or 20 (FR20) pecks on the sample stimulus. None of the birds showed any evidence of concept-mediated transfer. In Experiment 2, 12 pigeons were trained with 3 stimuli and then tested with the same novel stimuli used in Experiment 1. Half of the birds in each group were trained with either an FR1 or FR20 requirement on the sample stimulus. Two of the FR20 birds showed high levels of transfer to the novel stimuli similar to that of monkeys in a previous study.     

Internal sense of direction and landmark use in pigeons (Columba livia)

The relative importance of an internal sense of direction based on inertial cues and landmark piloting for small-scale navigation by White King pigeons (Columba livia) was investigated in an arena search task. Two groups of pigeons differed in whether they had access to visual cues outside the arena. In Experiment 1, pigeons were given experience with 2 different entrances and all pigeons transferred accurate searching to novel entrances. Explicit disorientation before entering did not affect accuracy. In Experiments 2-4, landmarks and inertial cues were put in conflict or tested 1 at a time. Pigeons tended to follow the landmarks in a conflict situation but could use an internal sense of direction to search when landmarks were unavailable.     

Perceptual completion in rhesus monkeys (Macaca mulatta) and pigeons (Columba livia)

In a two-dimensional drawing, when the narrow edge of a bar appears to touch the edge of a large rectangle, humans overestimate the length of the bar (Kanizsa, 1979). Kanizsa has suggested that this illusion occurs because humans perceive the bar as continuing behind the rectangle and complete the “occluded” portion of the bar. Rhesus monkeys and pigeons were trained to classify black target bars with a variety of lengths as “long” or “short.” In training, the bar was always located at the same distance from a gray box. After learning this discrimination, the subjects were tested on novel stimuli, in which the bar was located at three new locations. Monkeys showed a consistent response bias for “long” when the bar touched the box, but pigeons did not. Monkeys appear to have completed the “occluded” part like humans, whereas pigeons failed to do so. Because this procedure does not require animals to complete the “occluded” part with any particular form, their failure suggests that pigeons do not even perceive the target bar as continuing behind the “occluding” figure. The failure of pigeons may be due to difficulty in perceiving depth from two-dimensional drawings.     

Septal area lesions impair spatial working memory in homing pigeons (Columba livia)

The septo-hippocampal system in birds resembles that of mammals, motivating research into the function of the avian hippocampus while surprisingly little attention has been given to the septum. To investigate a possible role of the avian septum in memory, the effects of septal area lesions on a spatial working memory (SpWM) task was tested in homing pigeons. After preoperative training on an analogue eight-arm (feeders) radial maze, now sham-operated control and septal lesioned pigeons were then trained again on the same task of locating the four feeders on the test phase of a trial that were not baited during the sample phase of a trial. During the test phase of a working memory trial, septal lesioned pigeons, compared to both their own preoperative performance and the performance of the controls, required significantly more choices to locate the four baited feeders not baited during the sample phase of a trial, and they made significantly fewer correct responses to the now baited feeders on their first four choices. The results demonstrate that, like its mammalian counterpart, the avian septum plays an important role in SpWM, suggesting that at least some functional properties of the septum are evolutionarily conserved in birds and mammals.     

Qualities and locations of stimuli and responses affecting discrimination learning of chinchillas (Chinchilla laniger) and pigeons (Columba livia)

Chinchillas and pigeons were used as subjects in separate experiments to study interactions among stimulus and response characteristics in discrimination learning. Both the stimuli and the responses could differ with respect to their "quality" and their "location." Bright versus dim lights and upper versus lower lights served as the stimulus qualities and stimulus locations for the chinchillas, respectively. Red versus green lights and upper versus lower lights served as the stimulus qualities and stimulus locations for the pigeons, respectively. Respond versus no-respond and respond-left versus respond-right served as the response qualities and response locations, for both species, respectively. In both experiments, response-quality performance was superior when the discriminative stimuli differed in quality than when they differed in location, whereas response-location performance was superior when the discriminative stimuli differed in location than when they differed in quality. These results were interpreted within the framework provided by a general law of learning, that is, the "quality-location hypothesis."     

Failure of hierarchical conditional rule learning in the pigeon (Columba livia)

Pigeons were trained with a conditional discrimination task in three-key operant chambers. Choosing either the left or right key was followed by food according to combinations of three preceding events: (a) a houselight illumination condition (dark or light), (b) presence or absence of green flashes on the three keys, (c) a color (amber or blue) of the center sample key. With these 2×2×2 event combinations, eight types of correct trials were prepared: (1) dark→no flash→amber→LEFT, (2) dark→no flash→blue→RIGHT, (3) dark→flash→amber→RIGHT, (4) dark→flash→blue→LEFT, (5) light→no flash→amber→RIGHT, (6) light→no flash→blue→LEFT, (7) light→flash→amber→LEFT, and (8) light→flash→blue→RIGHT. Seven of these eight types were used for training of a given bird, and then the remaining trial type was presented as the test. If the birds had learned the conditional structure of the events (the hierarchical switching rule), they would have responded correctly to the test type. However, they chose the opposite side key, suggesting that they had learned cue configuration or multiple rules to solve the task. Accepted after revision: 13 January 2001 Electronic Publication     

Symmetry and transitivity of conditional relations in monkeys (Cebus apella) and pigeons (Columba livia)

In Experiment 1 six monkeys were tested with discriminative relations that were backward relative to their training in a 0-second conditional (“symbolic”) matching procedure. Although there was some indication of backward associations, the evidence was generally weak, and statistical evaluations did not reach conventional significance levels. Unlike children, who show backward associations to the point of symmetry, monkeys and pigeons display at best only weak and transient backward associations. In Experiment 2 associative transitivity was assessed across two sets of conditional matching tasks. All four monkeys tested demonstrated strong transitivity. In contrast, in Experiment 3 there was no evidence of transitivity in three pigeons tested under conditions closely comparable to those of Experiment 2. These results may identify some key features of interspecies differences and contribute to analyses of serial learning in animals.     

Hippocampal lesions in pigeons (Columba livia) disrupt reinforced preexposure but not overshadowing or blocking

Three experiments examined the effects of hippocampal lesions in pigeons on overshadowing, blocking, and a latent inhibition treatment (non-differential reinforced preexposure) using a simultaneous visual discrimination paradigm. The results showed that hippocampal damage did not influence overshadowing (Experiment 1) or blocking (Experiment 2) but did attenuate the retardation in conditioning associated with non-differential reinforced preexposure to to-be-discriminated stimuli (Experiment 3). Hippocampal birds also displayed impaired autoshaping (Experiments 1, 2, and 3). The correspondence between the behavioural effects of avian and mammalian hippocampal lesions is discussed, and the implications of the present pattern of results for avian hippocampal function are considered.