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Petruso, E. J., Fuchs, T., & Bingman, V. P. (2007). Time-space learning in homing pigeons (Columba livia): orientation to an artificial light source. Anim. Cogn., 10(2), 181–188.
Abstract: Time-space learning reflects an ability to represent in memory event-stimulus properties together with the place and time of the event; a capacity well developed in birds. Homing pigeons were trained in an indoor octagonal arena to locate one food goal in the morning and a different food goal in the late afternoon. The goals differed with respect to their angular/directional relationship to an artificial light source located outside the arena. Further, the angular difference in reward position approximated the displacement of the sun's azimuth that would occur during the same time period. The experimental birds quickly learned the task, demonstrating the apparent ease with which birds can adopt an artificial light source to discriminate among alternative spatial responses at different times of the day. However, a novel midday probe session following successful learning revealed that the light source was interpreted as a stable landmark and not as a surrogate sun that would support compass orientation. Probe sessions following a phase shift of the light-dark cycle revealed that the mechanism employed to make the temporal discrimination was prevailingly based on an endogenous circadian rhythm and not an interval timing mechanism.
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Fairhurst, S., Gallistel, C. R., & Gibbon, J. (2003). Temporal landmarks: proximity prevails. Anim. Cogn., 6(2), 113–120.
Abstract: Subjects in conditioning experiments time their conditioned responses relative to the onsets of the conditioned stimuli (CSs). These onsets are temporal landmarks, by reference to which subjects may estimate the location of the unconditioned stimulus (US) in time. In a serial compound conditioning paradigm, a long duration CS comes on first, followed later by a second shorter CS, creating both a long-range and a short-range predictor of the US. We ask whether displacing the short-range predictor relative to the long-range predictor causes subjects to strike a compromise between the different temporal locations predicted by the two CSs. In three experiments with pigeons, we varied the training conditions so as to favor or militate against this outcome. However, in all conditions, there was no compromise; after the onset of the displaced short-range CS, the timing of conditioned responding was governed by it alone. This result contrasts with the compromises that are seen when the feeding time predicted by a CS is put in conflict with the time predicted by the circadian clock, and with the similar compromises sometimes seen when a nearby spatial landmark is displaced relative to a larger spatial context.
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Cerutti, D. T., & Staddon, J. E. R. (2004). Immediacy versus anticipated delay in the time-left experiment: a test of the cognitive hypothesis. J Exp Psychol Anim Behav Process, 30(1), 45–57.
Abstract: In the time-left experiment (J. Gibbon & R. M. Church, 1981), animals are said to compare an expectation of a fixed delay to food, for one choice, with a decreasing delay expectation for the other, mentally representing both upcoming time to food and the difference between current time and upcoming time (the cognitive hypothesis). The results of 2 experiments support a simpler view: that animals choose according to the immediacies of reinforcement for each response at a time signaled by available time markers (the temporal control hypothesis). It is not necessary to assume that animals can either represent or subtract representations of times to food to explain the results of the time-left experiment.
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Crystal, J. D. (1999). Systematic nonlinearities in the perception of temporal intervals. J Exp Psychol Anim Behav Process, 25(1), 3–17.
Abstract: Rats judged time intervals in a choice procedure in which accuracy was maintained at approximately 75% correct. Sensitivity to time (d') was approximately constant for short durations 2.0-32.0 s with 1.0- or 2.0-s spacing between intervals (n = 5 in each group, Experiment 1), 2.0-50.0 s with 2.0-s spacing (n = 2, Experiment 1), and 0.1-2.0 s with 0.1- or 0.2-s spacing (n = 6 in each group, Experiment 2). However, systematic departures from average sensitivity were observed, with local maxima in sensitivity at approximately 0.3, 1.2, 10.0, 24.0, and 36.0 s. Such systematic departures from an approximately constant d' are predicted by a connectionist theory of time with multiple oscillators and may require a modification of the linear timing hypothesis of scalar timing theory.
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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., & Kaiser, D. H. (2005). Interval timing with gaps: gap ambiguity as an alternative to temporal decay. J Exp Psychol Anim Behav Process, 31(4), 484–486.
Abstract: C. V. Buhusi, D. Perera, and W. H. Meck (2005) proposed a hypothesis of timing in rats to account for the results of experiments that have used the peak procedure with gaps. According to this hypothesis, the introduction of a gap causes the animal's memory for the pregap interval to passively decay (subjectively shorten) in direct proportion to the duration and salience of the gap. Thus, animals should pause with short, nonsalient gaps but should reset their clock with longer, salient gaps. The present authors suggest that the ambiguity of the gap (i.e., the similarity between the gap and the intertrial interval in both appearance and relative duration) causes the animal to actively reset the clock and prevents adequate assessments of the fate of timed intervals prior to the gap. Furthermore, when the intertrial interval is discriminable from the gap, the evidence suggests that timed intervals prior to the gap are not lost but are retained in memory.
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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., Weaver, J. E., & Clement, T. S. (2004). Pigeons group time intervals according to their relative duration. Psychon Bull Rev, 11(1), 113–117.
Abstract: In the present research, we asked whether pigeons tended to judge time intervals not only in terms of their absolute value but also relative to a duration from which they must be discriminated (i.e., longer or shorter). Pigeons were trained on two independent temporal discriminations. In one discrimination, sample durations of 2 and 8 sec were associated with, for example, red and green hue comparisons, respectively, and in the other discrimination, sample durations of 4 and 16 sec were associated with vertical and horizontal line comparisons, respectively. If pigeons are trained on a temporal discrimination and tested with intermediate durations, the subjective midpoint typically occurs close to the geometric mean of the two trained values. The 4- and 8-sec values were selected to be the geometric mean of the two values in the other discrimination. When a 4-sec test sample was presented with the comparisons from the 2- and 8-sec discrimination, the pigeons preferred the comparison associated with the shorter sample. Similarly, when an 8-sec test sample was presented with the comparisons from the 4- and 16-sec discrimination, the pigeons preferred the comparison associated with the longer sample. Thus, a relative grouping effect was found. That is, durations that should have produced indifferent choice were influenced by their relative durations (shorter than or longer than the alternative) during training.
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Zentall, T. R., Klein, E. D., & Singer, R. A. (2004). Evidence for detection of one duration sample and default responding to other duration samples by pigeons may result from an artifact of retention-test ambiguity. J Exp Psychol Anim Behav Process, 30(2), 129–134.
Abstract: S. C. Gaitan and J. T. Wixted (2000) proposed that when pigeons are trained on a conditional discrimination to associate 1 duration sample with 1 comparison and 2 other duration samples with a 2nd comparison, they detect only the single duration, and on trials involving either of the 2 other duration samples, they respond to the other comparison by default. In 2 experiments, the authors show instead that pigeons lend to treat the retention intervals (such as those used by Gaitan and Wixted) as intertrial intervals, and thus, they tend to treat all trials with a delay as 0-s sample trials. The authors tested this hypothesis by showing that divergent retention functions do not appear when the retention interval is discriminably different from the intertrial interval.
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Kaiser, D. H., Zentall, T. R., & Neiman, E. (2002). Timing in pigeons: effects of the similarity between intertrial interval and gap in a timing signal. J Exp Psychol Anim Behav Process, 28(4), 416–422.
Abstract: Previous research suggests that when a fixed interval is interrupted (known as the gap procedure), pigeons tend to reset memory and start timing from 0 after the gap. However, because the ambient conditions of the gap typically have been the same as during the intertrial interval (ITI), ambiguity may have resulted. In the present experiment, the authors found that when ambient conditions during the gap were similar to the ITI, pigeons tended to reset memory, but when ambient conditions during the gap were different from the ITI, pigeons tended to stop timing, retain the duration of the stimulus in memory, and add to that time when the stimulus reappeared. Thus, when the gap was unambiguous, pigeons timed accurately.
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