Attention in the pigeon: a reevaluation

During training sessions, pigeons were successively exposed to compounds consisting of a white triangle on a red background and a white circle on a green background. Key pecking intermittently produced grain reinforcers in the presence of one form-color compound. Once key pecking was confined to the compound associated with reinforcement, the elements—red, green, triangle, and circle—were presented during a test in which no reinforcement was available. Each bird pecked nearly exclusively in the presence of the color previously associated with reinforcement, a result that might be interpreted as indicating that the subjects had attended to color, but not form during training. Pecking was next reinforced when either the triangle or the circle was present. Pecking in the presence of the form previously associated with reinforcement was acquired more rapidly. This result suggests that the birds had learned about the forms during training, and that conclusions about attention based on the lack of differential pecking in the nonreinforcement test may not be appropriate.     

Detecting a nonevent: Delayed presence-versus-absence discrimination in pigeons

Eight pigeons were trained on a delayed presence-versus-absence discrimination paradigm in which a sample stimulus was presented on some trials but not on others. If a sample was presented, then a response to one choice key produced food. If no sample was presented, a response to the other choice key produced food. The basic finding was that performance remained constant and well above 50% correct on no-sample trials as the retention interval increased, whereas performance dropped precipitously (to below 50% correct) on sample trials. In the second phase of the experiment, all of the trials were no-sample trials, and reinforcers were delivered probabilistically for one group of pigeons and according to time-based schedules for the other group. The exact reinforcement probabilities used in Phase 2 were those calculated to be in effect on no-sample trials in Phase 1 (according to a discrete-state model of performance). Subjects did not show exclusive preference for the richer alternative on no-sample trials in the first phase, but those in the probabilistic group developed near-exclusive preference for the richer alternative during the second phase. These data are inconsistent with the predictions of the discrete-state model, but are easily accommodated by an account based on signal detection theory, which also can be applied effectively to discrimination of event duration and the “subjective shortening” effect.     

Two routes toward optimism: how agentic and communal themes in autobiographical memories guide optimism for the future

Autobiographical memories are particularly adaptive because they function not only to preserve the past, but also to direct our future thoughts and behaviours. Two studies were conducted to examine how communal and agentic themes of positive autobiographical memories differentially predicted the route from autobiographical memories to optimism for the future. Across two studies, results revealed that the degree to which participants focused on communal themes in their autobiographical memories predicted their experience of nostalgia. In turn, the experience of nostalgia increased participants’ levels of self-esteem and in turn, optimism for the future. By contrast, the degree to which participants focused on agentic themes in their memories predicted self-esteem and optimism, operating outside the experience of nostalgia. These effects remained even after controlling for self-focused attention. Together, these studies provide greater understanding of the interrelations among autobiographical memory, self-concept, and time, and demonstrate how agency and communion operate to influence perceptions of one’s future when thinking about the past.     

Bonobos and orangutans,but not chimpanzees,flexibly plan for the future in a token-exchange task

Non-human animals, including great apes, have been suggested to share some of the skills for planning that humans commonly exhibit. A crucial difference between human and non-human planning may relate to the diversity of domains and needs in which this skill is expressed. Although great apes can save tools for future use, there is little evidence yet that they can also do so in other contexts. To investigate this question further, we presented the apes with a planning token-exchange task that differed from standard tool-use tasks. Additionally, we manipulated the future outcome of the task to investigate planning flexibility. In the Exchange condition, subjects had to collect, save and transport tokens because they would need them 30 min later to exchange them for food with a human, i.e., “bring-back” response. In the Release condition, the collection and transport of tokens were not needed as no exchange took place after 30 min. Out of 13 subjects, eight solved the task at least once in the Exchange condition, with chimpanzees appearing less successful than the other species. Importantly, three individuals showed a clear differential response between conditions by producing more “bring-back” responses in the Exchange than in the Release conditions. Those bonobo and orangutan individuals hence adapted their planning behavior according to changing needs (i.e., they brought tokens back significantly more often when they would need them). Bonobos and orangutans, unlike chimpanzees, planned outside the context of tool-use, thus challenging the idea that planning in these species is purely domain-specific.     

Social complexity can drive vocal complexity: group size influences vocal information in Carolina chickadees

One hypothesis to explain variation in vocal communication in animal species is that the complexity of the social group influences the group's vocal complexity. This social-complexity hypothesis for communication is also central to recent arguments regarding the origins of human language, but experimental tests of the hypothesis are lacking. This study investigated whether group size, a fundamental component of social complexity, influences the complexity of a call functioning in the social organization of Carolina chickadees, Poecile carolinensis. In unmanipulated field settings, calls of individuals in larger groups had greater complexity (more information) than calls of individuals in smaller groups. In aviary settings manipulating group size, individuals in larger groups used calls with greater complexity than individuals in smaller groups. These results indicate that social complexity can influence communicative complexity in this species.     

Autoshaping as a psychophysical paradigm: Absolute visual sensitivity in the pigeon

A classical conditioning procedure (autoshaping) was used to determine absolute visual threshold in the pigeon. This method provides the basis for a standardized visual psychophysical paradigm.     

Delay discounting in the pigeon: In search of a magnitude effect

The magnitude effect, where larger outcomes are discounted proportionally less than smaller outcomes, is a well‐established phenomenon in delay discounting by human participants. To this point in the literature magnitude effects have not been reliably evidenced in nonhuman animals. , however, used a concurrent‐chains arrangement with pigeon and found evidence for a magnitude effect. Grace et al. suggested that in many delay discounting experimental arrangements with nonhuman animals (e.g., adjusting amount, adjusting delay) the organism is not given the opportunity to directly compare outcomes of different sizes. They suggest that because of the lack of direct comparison it is difficult for the organism to determine the relative size of each outcome, which in turn mutes the effect of the amount differences between outcomes. As a test of this “comparison hypothesis,” the present experiment was conducted to assess whether the magnitude effect would be evidenced in pigeon when using an adjusting amount procedure where outcomes of different amounts were presented proximally. In the present arrangement, pigeons were presented two choice panels in an operant chamber where each panel was associated with an independent adjusting amount delay discounting task, but with differing outcome amounts (i.e., a 32‐food pellet panel and an 8‐food pellet panel). In this arrangement the choice panels alternated in their availability within a session from trial block to trial block. The present findings indicate no reliable effect of amount, even when the outcomes were proximal and thus readily comparable. This result suggests that the lack of magnitude effect is not driven by the organism's ability to compare the difference in amount between choice alternatives.     

Successive matching-to-sample in the pigeon: Variations on a theme by Konorski

In 1959, Konorski proposed a successive matching-to-sample paradigm with which to study short-term memory in animals. Although the technique has been little used, it affords several distinct advantages over more commonly employed matching-to-sample procedures. Konorski’s paradigm involves the successive presentation of a pair of discriminative stimuli with a brief interstimulus interval between them. Reinforcement is scheduled to occur only when the second stimulus of a pair matches the first; otherwise, nonreinforcement follows. An investigation of the pigeon’s food-reinforced keypecking behavior is reported using a variant of Konorski’s technique. Pigeons rapidly learned to differentiate matching and nonmatching stimulus pairs when brief (5-sec) color stimuli were separated by a 1-sec interstimulus interval. No such differentiation arose when control subjects were trained with reinforcement equiprobable on matching and nonmatching trials. No support was found for the notion that correct performance under this successive-matching procedure was due to overt mediating behaviors.     

Figure-ground assignment in pigeons: evidence for a figural benefit

Four pigeons discriminated whether a target spot appeared on a colored figural shape or on a differently colored background by first pecking the target and then reporting its location: on the figure or the background. We recorded three dependent variables: target detection time, choice response time, and choice accuracy. The birds were faster to detect the target, to report its location, and to learn the correct response on figure trials than on background trials. Later tests suggested that the pigeons might have attended to the figural region as a whole rather than using local properties in performing the figure-background discrimination. The location of the figural region did not affect figure-ground assignment. Finally, when 4 other pigeons had to detect and peck the target without making a choice report, no figural advantage emerged in target detection time, suggesting that the birds' attention may not have been automatically summoned to the figural region.     

Short-term memory in the pigeon: stimulus-response associations

Three pigeons pecked for food in two experiments in which each trial consisted of two phases: a study and a test phase. The study phase in Experiment I consisted of two stimulus-response pairs presented successively. Each pair consisted of the illumination of a left or right key (the stimulus) and a peck on the lighted side key (the response). The study phase in Experiment II consisted of three such pairs presented successively. A retention interval, varied between 0.1 and 4.0 sec, separated the study phase from the test phase. The test phase of a trial began with the illumination of the center key by one of two (Experiment I) or three (Experiment II) colors. This color was the same as the stimulus element of one of the pairs in the study phase. A reinforcer was presented if a subject then emitted the response element of the indicated stimulus-response pair. The results provide information on the conditions that enable a pigeon to remember the responses most recently emitted in the presence of various stimuli. The results suggest an account of the maintenance of behavior that is temporally noncontiguous with reinforcement.     

Short-term memory in the pigeon: relative recency

Three pigeons pecked for food in an experiment in which each trial consisted of two phases. The first phase consisted of a pattern of three successively illuminated, randomly selected left or right keys. A subject was required to peck each of the lighted keys as they appeared. Thus, in the first phase, a subject emitted a pattern of three left- or right-key pecks. Over trials, all eight possible patterns appeared. A time interval separated the first phase from the second phase, which began with presentation of a randomly selected one of three cues. A reinforcer was delivered in the second phase if a subject pecked the side key that had appeared in the first phase in an ordinal position corresponding to the cue presented in the second phase. That is, the three cues probed a pigeon's memory for the side key it had pecked first, second, or third, in the first phase of a trial. The results show that a pigeon can remember for more than 4 sec the order in which it has just seen and pecked two lighted keys: a pigeon can remember the temporal organization or pattern of events in its recent environment. Consequently, the functional stimulus present when a reinforcer is delivered may include a subject's short-term memory for the temporal organization of recent events, such as the pattern of its own recent behavior. This possibility is consistent with a molecular analysis of operant behavior focusing on local patterns of behavior.     

Beacon training in a water maze can facilitate and compete with subsequent room cue learning in rats

In Stage 1 of 4 experiments in which rats completed a water-maze blocking procedure, experimental groups were trained to use a predictive beacon (hanging above, connected to, or displaced from the platform) to find a submerged escape platform in the presence of predictive or irrelevant background cues and in the presence or absence of irrelevant landmarks. In Stage 2, a fixed beacon, landmarks, and background cues all predicted the platform location. A Room Test (landmarks and background cues only) showed that Stage 1 training with a fixed hanging beacon or the moving displaced beacon facilitated Stage 2 learning of predictive room cues for experimental relative to control subjects. In contrast, Stage 1 training with a moving pole beacon interfered with Stage 2 learning about predictive room cues relative to controls, whereas training with a fixed pole or moving hanging beacon had no effect. We conclude that multiple spatial learning processes influence locating an escape platform in the water maze.     

Free-operant avoidance in the pigeon using a treadle response

A free-operant avoidance schedule was used to establish and maintain foot-treadle responding by two Homing, one White King, and two Carneaux pigeons. In the absence of responding, the interval between shocks equaled 10 sec. Each time a treadle response occurred the shock was postponed for 32 sec. Pigeons appear to learn the treadle response more quickly and use it to avoid shock more successfully than do rats bar pressing on similar schedules. The treadle response becomes highly stereotyped and interresponse time distributions obtained from terminal behavior appear very similar to data obtained from rats. It is concluded that the difficulty in training pigeons to avoid electric shock is not in establishing avoidance behavior but in attempting to evaluate such behavior with the key-peck response.     

Short-term memory in the pigeon: the previously reinforced response

Eighteen pigeons served in a discrete-trials short-term memory experiment in which the reinforcement probability for a peck on one of two keys depended on the response reinforced on the previous trial: either the probability of reinforcement on a trial was 0.8 for the same response reinforced on the previous trial and was 0.2 for the other response (Group A), or, it was 0 or 0.2 for the same response and 1.0 or 0.8 for the other response (Group B). A correction procedure ensured that over all trials reinforcement was distributed equally across the left and right keys. The optimal strategy was either a winstay, lose-shift strategy (Group A) or a win-shift, lose-stay strategy (Group B). The retention interval, that is the intertrial interval, was varied. The average probability of choosing the optimal alternative reinforced 80% of the time was 0.96, 0.84, and 0.74 after delays of 2.5, 4.0, and 6.0 sec, respectively for Group A, and was 0.87, 0.81, and 0.55 after delays of 2.5, 4.0, and 6.0 sec, respectively, for Group B. This outcome is consistent with the view that behavior approximated the optimal response strategy but only to an extent permitted by a subject's short-term memory for the cue correlated with reinforcement, that is, its own most-recently reinforced response. More generally, this result is consistent with “molecular” analyses of operant behavior, but is inconsistent with traditional “molar” analyses holding that fundamental controlling relations may be discovered by routinely averaging over different local reinforcement contingencies. In the present experiment, the molar results were byproducts of local reinforcement contingencies involving an organism's own recent behavior.     

Food delivery as a conditional stimulus: Feature-learning and memory in pigeons

Three experiments investigated the learning and memory of discriminations based on presence versus absence of a pre-trial food delivery. In Experiment 1 half the illuminations of a response key were followed by food regardless of the subject's behavior. In one group an extra food delivery preceded only reinforced trials (feature-positive condition), whereas in a second group it preceded only nonreinforced trials (feature-negative condition). Key pecks and approaches revealed more rapid and superior discrimination learning in the first group. Experiment 2 replicated the results of Experiment 1 but yielded no evidence that greater “unexpectedness” of pretrial food conditions facilitates discriminative performance. In Experiment 3, individual pigeons trained on a conditional discrimination exhibited a within-subject feature-positive superiority. Delay between pretrial and trial stimuli interacted with feature-positive versus feature-negative training in both the between-group (Experiment 2) and within-subject (Experiment 3) procedures: performance was decremented at both short and long delays in the feature-positive condition but was decremented only at longer delays in the feature-negative condition. The feature-positive superiority obtained here is incompatible with explanations based on either the general concept of “perceptual organization” or on the conditional nature of feature-negative discriminations.     

Categories in the pigeon brain: A reverse engineering approach

Pigeons are well known for their visual capabilities as well as their ability to categorize visual stimuli at both the basic and superordinate level. We adopt a reverse engineering approach to study categorization learning: Instead of training pigeons on predefined categories, we simply present stimuli and analyze neural output in search of categorical clustering on a solely neural level. We presented artificial stimuli, pictorial and grating stimuli, to pigeons without the need of any differential behavioral responding while recording from the nidopallium frontolaterale (NFL), a higher visual area in the avian brain. The pictorial stimuli differed in color and shape; the gratings differed in spatial frequency and amplitude. We computed representational dissimilarity matrices to reveal categorical clustering based on both neural data and pecking behavior. Based on neural output of the NFL, pictorial and grating stimuli were differentially represented in the brain. Pecking behavior showed a similar pattern, but to a lesser extent. A further subclustering within pictorial stimuli according to color and shape, and within gratings according to frequency and amplitude, was not present. Our study gives proof‐of‐concept that this reverse engineering approach—namely reading out categorical information from neural data—can be quite helpful in understanding the neural underpinnings of categorization learning.     

Auto-maintenance in the pigeon: sustained pecking despite contingent non-reinforcement

If a response key is regularly illuminated for several seconds before food is presented, pigeons will peck it after a moderate number of pairings; this “auto-shaping” procedure of Brown and Jenkins (1968) was explored further in the present series of four experiments. The first showed that pecking was maintained even when pecks turned off the key and prevented reinforcement (auto-maintenance); the second controlled for possible effects of generalization and stimulus change. Two other experiments explored procedures that manipulated the tendency to peck the negatively correlated key by introducing alternative response keys which had no scheduled consequences. The results indicate that pecking can be established and maintained by certain stimulus-reinforcer relationships, independent of explicit or adventitious contingencies between response and reinforcer.     

Interocular generalization: a study of mirror-image reversal following monocular discrimination training in the pigeon

Generalization gradients along a continuum of angular orientation were obtained from four pigeons, following monocular training on a discrimination between a 45° oblique line (S+) and a 135° oblique line (S−). All pigeons were trained on a chain DRO VI 1 schedule of reinforcement. Generalization gradients obtained with the trained and untrained eye were compared. All pigeons responded maximally to the 45° line (S+) when tested with only the trained eye open. During generalization tests of interocular transfer with only the untrained eye open, three pigeons responded maximally to S− (135°), the mirror-image of the stimulus associated with reinforcement during training (45°). The other pigeon failed to show interocular transfer of the discrimination. Interocular reversal of left-right mirror-image stimuli has not been reported for any other species.