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Zentall, T. R., Clement, T. S., Bhatt, R. S., & Allen, J. (2001). Episodic-like memory in pigeons. Psychon Bull Rev, 8(4), 685–690.
Abstract: It has been proposed that memory for personal experiences (episodic memory, rather than semantic memory) relies on the conscious review of past experience and thus is unique to humans. In an attempt to demonstrate episodic-like memory in animals, we first trained pigeons to respond to the (nonverbal) question “Did you just peck or did you just refrain from pecking?” by training them on a symbolic matching task with differential responding required to the two line-orientation samples and reinforcing the choice of a red comparison if they had pecked and the choice of a green comparison if they had not pecked. Then, in Experiment 1, after providing the conditions for (but not requiring) the pigeons to peck at one new stimulus (a yellow hue) but not at another (a blue hue), we tested them with the new hue stimuli and the red and green comparisons. In Experiment 2, we tested the pigeons with novel stimuli (a circle, which they spontaneously pecked, and a dark response key, which they did not peck) and the red and green comparisons. In both experiments, pigeons chose the comparison appropriate to the response made to the test stimulus. Thus, the pigeons demonstrated that they could remember specific details about their past experiences, a result consistent with the notion that they have the capacity for forming episodic-like memories.
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Zentall, T. R., & Clement, T. S. (2002). Memory mechanisms in pigeons: evidence of base-rate neglect. J Exp Psychol Anim Behav Process, 28(1), 111–115.
Abstract: In delayed matching to sample, once acquired, pigeons presumably choose comparisons according to their memory for (the strength of) the sample. When memory for the sample is sufficiently weak, comparison choice should depend on the history of reinforcement associated with each of the comparison stimuli. In the present research, pigeons acquired two matching tasks in which Sample S1 was associated with one comparison from each task, C1 and C3, whereas Sample S2 was associated with Comparison C2, and Sample S3 was associated with Comparison C4. As the retention interval increased, the pigeons showed a bias to choose the comparison (C1 or C3) associated with the more frequently occurring sample (S1). Thus, pigeons were sensitive also to the (irrelevant) likelihood that each of the samples was presented. The results suggest that pigeons may allow their reference memory for the overall sample frequency to influence comparison choice, independent of the comparison stimuli present.
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Zentall, T. R. (2006). Mental time travel in animals: a challenging question. Behav. Process., 72(2), 173–183.
Abstract: Humans have the ability to mentally recreate past events (using episodic memory) and imagine future events (by planning). The best evidence for such mental time travel is personal and thus subjective. For this reason, it is particularly difficult to study such behavior in animals. There is some indirect evidence, however, that animals have both episodic memory and the ability to plan for the future. When unexpectedly asked to do so, animals can report about their recent past experiences (episodic memory) and they also appear to be able to use the anticipation of a future event as the basis for a present action (planning). Thus, the ability to imagine past and future events may not be uniquely human.
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Zentall, T. R. (2005). Timing, memory for intervals, and memory for untimed stimuli: the role of instructional ambiguity. Behav. Process., 70(3), 209–222.
Abstract: Theories of animal timing have had to account for findings that the memory for the duration of a timed interval appears to be dramatically shorted within a short time of its termination. This finding has led to the subjective shortening hypothesis and it has been proposed to account for the poor memory that animals appear to have for the initial portion of a timed interval when a gap is inserted in the to-be-timed signal. It has also been proposed to account for the poor memory for a relatively long interval that has been discriminated from a shorter interval. I suggest here a simpler account in which ambiguity between the gap or retention interval and the intertrial interval results in resetting the clock, rather than forgetting the interval. The ambiguity hypothesis, together with a signal salience mechanism that determines how quickly the clock is reset at the start of the intertrial interval can account for the results of the reported timing experiments that have used the peak procedure. Furthermore, instructional ambiguity rather than memory loss may account for the results of many animal memory experiments that do not involve memory for time.
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Zentall, T. R. (1999). Support for a theory of memory for event duration must distinguish between test-trial ambiguity and actual memory loss. J Exp Anal Behav, 72(3), 467–472.
Abstract: Staddon and Higa's (1999) trace-strength theory of timing and memory for event duration can account for pigeons' bias to “choose short” when retention intervals are introduced and to “choose long” when, following training with a fixed retention interval, retention intervals are shortened. However, it does not account for the failure of pigeons to choose short when the intertrial interval is distinct from the retention interval. That finding suggests that stimulus generalization (or ambiguity) between the intertrial interval and the retention interval may result in an effect that has been attributed to memory loss. Such artifacts must be eliminated before a theory of memory for event duration can be adequately tested.
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Watanabe, S., & Troje, N. F. (2006). Towards a “virtual pigeon”: a new technique for investigating avian social perception. Anim. Cogn., 9(4), 271–279.
Abstract: The purpose of the present study is to examine the applicability of a computer-generated, virtual animal to study animal cognition. Pigeons were trained to discriminate between movies of a real pigeon and a rat. Then, they were tested with movies of the computer-generated (CG) pigeon. Subjects showed generalization to the CG pigeon, however, they also responded to modified versions in which the CG pigeon was showing impossible movement, namely hopping and walking without its head bobbing. Hence, the pigeons did not attend to these particular details of the display. When they were trained to discriminate between the normal and the modified version of the CG pigeon, they were able to learn the discrimination. The results of an additional partial occlusion test suggest that the subjects used head movement as a cue for the usual vs. unusual CG pigeon discrimination.
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Wasserman, E. A., Gagliardi, J. L., Cook, B. R., Kirkpatrick-Steger, K., Astley, S. L., & Biederman, I. (1996). The pigeon's recognition of drawings of depth-rotated stimuli. J Exp Psychol Anim Behav Process, 22(2), 205–221.
Abstract: Four experiments used a four-choice discrimination learning paradigm to explore the pigeon's recognition of line drawings of four objects (an airplane, a chair, a desk lamp, and a flashlight) that were rotated in depth. The pigeons reliably generalized discriminative responding to pictorial stimuli over all untrained depth rotations, despite the bird's having been trained at only a single depth orientation. These generalization gradients closely resembled those found in prior research that used other stimulus dimensions. Increasing the number of different vantage points in the training set from one to three broadened the range of generalized testing performance, with wider spacing of the training orientations more effectively broadening generalized responding. Template and geon theories of visual recognition are applied to these empirical results.
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Wagner, G. (1975). [Flight leadership in flocks of homing pigeons]. Z. Tierpsychol., (39), 61–74.
Abstract: Groups of 3-5 homing pigeons individually recognizable by different colours of their plumage were followed by helicopter on their way home. In most cases the animals flew together as a group with frequently changing leadership. Flight formations in terms of leadership were noted every minute. It was examined statistically whether the flight order varies at random or whether there are leading and led birds. In 6 out of 7 experiments with groups of 4-5 pigeons flight order was far from random, one or two pigeons proving to be leaders. In only one experiment leadership did not differ from a random distribution. No correlation could be found between the tendency to lead within a group and homing performance of the single pigeon when released individually.
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Urcuioli, P. J., & Zentall, T. R. (1992). Transfer across delayed discriminations: evidence regarding the nature of prospective working memory. J Exp Psychol Anim Behav Process, 18(2), 154–173.
Abstract: Pigeons were trained successively either on 2 delayed simple discriminations or on a delayed simple discrimination followed by delayed matching-to-sample. During subsequent transfer tests, the initial stimuli from the 1st task were substituted for those in the 2nd. Performances transferred immediately if both sets of initial stimuli had been associated with the presence versus absence of food on their respective retention tests, and the direction of transfer (positive or negative) depended on whether the substitution involved stimuli with identical or different outcome associates. No transfer was found, however, when the initial stimuli were associated with different patterns of responding but food occurred at the end of every trial. These results are consistent with outcome expectancy mediation but are incompatible with response intention and retrospective coding accounts.
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Urcuioli, P. J., & Zentall, T. R. (1986). Retrospective coding in pigeons' delayed matching-to-sample. J Exp Psychol Anim Behav Process, 12(1), 69–77.
Abstract: In this study we examined how coding processes in pigeons' delayed matching-to-sample were affected by the stimuli to be remembered. In Experiment 1, two groups of pigeons initially learned 0-delay matching-to-sample with identical comparison stimuli (vertical and horizontal lines) but with different sample stimuli (red and green hues or vertical and horizontal lines). Longer delays were then introduced between sample offset and comparison onset to assess whether pigeons were prospectively coding the same events (viz., the correct line comparisons) or retrospectively coding different events (viz., their respective sample stimuli). The hue-sample group matched more accurately and showed a slower rate of forgetting than the line-sample group. In Experiment 2, pigeons were trained with either hues or lines as both sample and comparison stimuli, or with hue samples and line comparisons or vice versa. Subsequent delay tests revealed that the hue-sample groups remembered more accurately and generally showed slower rates of forgetting than the line-sample groups. Comparison dimension had little or no effect on performance. Together, these data suggest that pigeons retrospectively code the samples in delayed matching-to-sample.
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