Rebecca Rayburn-Reeves

Imitation and emulation by dogs using a bidirectional control procedure.

A successful procedure for studying imitative behavior in non-humans is the bidirectional control procedure in which observers are exposed to a demonstrator that responds by moving a manipulandum in one of two different directions (e.g., left vs. right). Imitative learning is demonstrated when observers make the response in the direction that they observed it being made. This procedure controls for socially mediated effects (the mere presence of a demonstrator), stimulus enhancement (attention drawn to a manipulandum by its movement), and if an appropriate control is included, emulation (learning how the environment works). Recent research with dogs has found that dogs may not demonstrate imitative learning when the demonstrator is human. In the present research, we found that when odors were controlled for, dogs imitated the direction of a screen-push demonstrated by another dog more than in a control condition in which they observed the screen move independently while another dog was present. Furthermore, we found that dogs would match the direction of screen-push demonstrated by a human and they were equally likely to match the direction in which the screen moved independently while a human was present.

Self-Control Without a “Self”? Common Self-Control Processes in Humans and Dogs

Self-control constitutes a fundamental aspect of human nature. Yet there is reason to believe that human and nonhuman self-control processes rely on the same biological mechanism—the availability of glucose in the bloodstream. Two experiments tested this hypothesis by examining the effect of available blood glucose on the ability of dogs to exert self-control. Experiment 1 showed that dogs that were required to exert self-control on an initial task persisted for a shorter time on a subsequent unsolvable task than did dogs that were not previously required to exert self-control. Experiment 2 demonstrated that providing dogs with a boost of glucose eliminated the negative effects of prior exertion of self-control on persistence; this finding parallels a similar effect in humans. These findings provide the first evidence that self-control relies on the same limited energy resource among humans and nonhumans. Our results have broad implications for the study of self-control processes in human and nonhuman species.

Object permanence in dogs: Invisible displacement in a rotation task

Dogs were tested for object permanence using an invisible displacement in which an object was hidden in one of two containers at either end of a beam and the beam was rotated. Consistent with earlier research, when the beam was rotated 180°, the dogs failed to find the object. However, when the beam was rotated only 90°, they were successful. Furthermore, when the dogs were led either 90° or 180° around the apparatus, they were also successful. In a control condition, when the dogs could not see the direction of the 90° rotation, they failed to find the object. The results suggest that the 180° rotation may produce an interfering context that can be reduced by rotating the apparatus only 90° or by changing the dogs’ perspective. Once the conflict is eliminated, dogs show evidence of object permanence that includes invisibly displaced objects.

Simultaneous discrimination reversal learning in pigeons and humans: anticipatory and perseverative errors

Pigeons were trained on a two-choice simultaneous discrimination (red vs. green) that reversed midway through each session. After considerable training, they consistently made both anticipatory errors prior to the reversal and perseverative errors after the reversal, suggesting that time (or number of trials) into the session served as a cue for reversal. In Experiment 2, to discourage the use of time as a cue, we varied the location of the reversal point within the session such that it occurred semirandomly after Trial 10, 25, 40, 55, or 70. Pigeons still tended both to anticipate and to perseverate. In Experiment 3, we required 20 pecks to a stimulus on each trial to facilitate memory for the preceding response and sensitivity to local reinforcement contingencies, but the results were similar to those of Experiment 2. We then tested humans on a similar task with a constant (Experiment 4) or variable (Experiment 5) reversal location. When the reversal occurred consistently at the midpoint of the session, humans, like pigeons, showed a tendency to anticipate the reversal; however, they did not show perseverative errors. When the reversal location varied between sessions, unlike pigeons, humans adopted a win–stay/lose–shift strategy, making only a single error on the first trial of the reversal.

Pigeons show near-optimal win-stay/lose-shift performance on a simultaneous-discrimination, midsession reversal task with short intertrial intervals.

Discrimination reversal tasks have been used as a measure of species flexibility in dealing with changes in reinforcement contingency. The simultaneous-discrimination, midsession reversal task is one in which one stimulus (S1) is correct for the first 40 trials of an 80-trial session and the other stimulus (S2) is correct for the remaining trials. After many sessions of training with this task, pigeons show a curious pattern of choices. They begin to respond to S2 well before the reversal point (they make anticipatory errors) and they continue to respond to S1 well after the reversal (they make perseverative errors). That is, they appear to be using the passage of time or number of trials into the session as a cue to reverse. We tested the hypothesis that these errors resulted in part from a memory deficit (the inability to remember over the intertrial interval, ITI, both the choice on the preceding trial and the outcome of that choice) by manipulating the duration of the ITI (1.5, 5, and 10 s). We found support for the hypothesis as pigeons with a short 1.5-s ITI showed close to optimal win-stay/lose-shift accuracy.

Midsession reversal learning: why do pigeons anticipate and perseverate?

Past research has shown that when given a simultaneous visual-discrimination midsession reversal task, pigeons typically anticipate the reversal well before it occurs and perseverate after it occurs. It appears that they use the estimation of time (or trial number) into the session, rather than (or in addition to) the more reliable cue, the outcome from the previous trial (i.e., a win–stay/lose–shift response rule), to determine which stimulus they should choose. In the present research, we investigated several variables that we thought might encourage pigeons to use a more efficient response strategy. In Experiment 1, we used a treadle-stepping response, rather than key pecking, to test the hypothesis that reflexive key pecking may have biased pigeons to estimate the time (or trial number) into the session at which the reversal would occur. In Experiment 2, we attempted to make the point of reversal in the session more salient by inserting irrelevant trials with stimuli different from the original discriminative stimuli, and for a separate group, we added a 5-s time-out penalty following incorrect choices. The use of a treadle-stepping response did not improve reversal performance, and although we found some improvement in reversal performance when the reversal was signaled and when errors resulted in a time-out, we found little evidence for performance that approached the win–stay/lose–shift accuracy shown by rats.

“counting” by pigeons: Discrimination of the number of biologically relevant sequential events

Numerical competence has been studied in animals under a variety of conditions, but only a few experiments have reported animals’ ability to detect absolute number. Capaldi and Miller (1988) tested rats’ ability to detect absolute number by using biologically important events—the number of reinforced runs followed by a nonreinforced run—and found that the rats ran significantly slower on the nonreinforced run. In the present experiments, we used a similar procedure. Pigeons were given a sequence of trials in which responding on the first three trials ended in reinforcement but responding on the fourth trial did not (RRRN). When the response requirement on each trial was a single peck (Experiment 1), we found no significant increase in latency to peck on the fourth trial. When the response requirement was increased to 10 pecks (Experiment 2), however, the time to complete the peck requirement was significantly longer on the nonreinforced trial than on the reinforced trials. Tests for control by time, number of responses, and amount of food consumed indicated that the pigeons were using primarily the number of reinforcements obtained in each sequence as a cue for nonreinforcement. This procedure represents a sensitive and efficient method for studying numerical competence in animals.

Who are the real bird brains? Qualitative differences in behavioral flexibility between dogs (Canis familiaris) and pigeons (Columba livia)

Pigeons given a simultaneous spatial discrimination reversal, in which a single reversal occurs at the midpoint of each session, consistently show anticipation prior to the reversal as well as perseveration after the reversal, suggesting that they use a less effective cue (time or trial number into the session) than what would be optimal to maximize reinforcement (local feedback from the most recent trials). In contrast, rats (Rattus norvegicus) and humans show near-optimal reversal learning on this task. To determine whether this is a general characteristic of mammals, in the present research, pigeons (Columba livia) and dogs (Canis familiaris) were tested with a simultaneous spatial discrimination mid-session reversal. Overall, dogs performed the task more poorly than pigeons. Interestingly, both pigeons and dogs employed what resembled a timing strategy. However, dogs showed greater perseverative errors, suggesting that they may have relatively poorer working memory and inhibitory control with this task. The greater efficiency shown by pigeons with this task suggests they are better able to time and use the feedback from their preceding choice as the basis of their future choice, highlighting what may be a qualitative difference between the species.

Pigeons’ use of cues in a repeated five-trial-sequence, single-reversal task

We studied behavioral flexibility, or the ability to modify one’s behavior in accordance with the changing environment, in pigeons using a reversal-learning paradigm. In two experiments, each session consisted of a series of five-trial sequences involving a simple simultaneous color discrimination in which a reversal could occur during each sequence. The ideal strategy would be to start each sequence with a choice of S1 (the first correct stimulus) until it was no longer correct, and then to switch to S2 (the second correct stimulus), thus utilizing cues provided by local reinforcement (feedback from the preceding trial). In both experiments, subjects showed little evidence of using local reinforcement cues, but instead used the mean probabilities of reinforcement for S1 and S2 on each trial within each sequence. That is, subjects showed remarkably similar behavior, regardless of where (or, in Exp. 2, whether) a reversal occurred during a given sequence. Therefore, subjects appeared to be relatively insensitive to the consequences of responses (local feedback) and were not able to maximize reinforcement. The fact that pigeons did not use the more optimal feedback afforded by recent reinforcement contingencies to maximize their reinforcement has implications for their use of flexible response strategies under reversal-learning conditions.

The Monty Hall dilemma in pigeons: Effect of investment in initial choice

In the Monty Hall dilemma, humans are initially given a choice among three alternatives, one of which has a hidden prize. After choosing, but before it is revealed whether they have won the prize, they are shown that one of the remaining alternatives does not have the prize. They are then asked whether they want to stay with their original choice or switch to the remaining alternative. Although switching results in obtaining the prize two thirds of the time, humans consistently fail to adopt the optimal strategy of switching even after considerable training. Interestingly, there is evidence that pigeons show more optimal switching performance with this task than humans. Because humans often view even random choices already made as being more valuable than choices not made, we reasoned that if pigeons made a greater investment, it might produce an endowment or ownership effect resulting in more human-like suboptimal performance. On the other hand, the greater investment in the initial choice by the pigeons might facilitate switching behavior by helping them to better discriminate their staying versus switching behavior. In Experiment 1, we examined the effect of requiring pigeons to make a greater investment in their initial choice (20 pecks rather than the usual 1 peck). We found that the increased response requirement facilitated acquisition of the switching response. In Experiment 2, we showed that facilitation of switching due to the increased response requirement did not result from extinction of responding to the initially chosen location.