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Shettleworth, S. J., & Krebs, J. R. (1982). How marsh tits find their hoards: the roles of site preference and spatial memory. J Exp Psychol Anim Behav Process, 8(4), 354–375.
Abstract: Marsh tits (Parus palustris) store single food items in scattered locations and recover them hours or days later. Some properties of the spatial memory involved were analyzed in two laboratory experiments. In the first, marsh tits were offered 97 sites for storing 12 seeds. They recovered a median of 65% of them 2-3 hr later, making only two errors per seed while doing so. Over trials, they used some sites more often than others, but during recovery they were more likely to visit a site of any preference value if they had stored a seed there that day than if they had not. Recovery performance was much worse if the experimenters moved the seeds between storage and recovery. A fixed search strategy that had some of the same average properties as the tits' search behavior also did worse than the real birds. In Experiment 2, any tendency to visit the same sites on successive daily tests in the aviary was placed in opposition to memory for storage sites by allowing the tits to store more seeds 2 hr after storing a first batch. They tended to avoid individual storage sites holding seeds from the first batch. When the tits searched for all the seeds 2 hr later, they tended to recover more seeds from the second batch than from the first, i.e., there was a recency effect.
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Shettleworth, S. J. (1985). Foraging, memory, and constraints on learning. Ann N Y Acad Sci, 443, 216–226.
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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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Rilling, M. E., & Neiworth, J. J. (1991). How animals use images. Sci Prog, 75(298 Pt 3-4), 439–452.
Abstract: Animal cognition is a field within experimental psychology in which cognitive processes formerly studied exclusively with people have been demonstrated in animals. Evidence for imagery in the pigeon emerges from the experiments described here. The pigeon's task was to discriminate, by pecking the appropriate choice key, between a clock hand presented on a video screen that rotated clockwise with constant velocity from a clock hand that violated constant velocity. Imagery was defined by trials on which the line rotated from 12.00 o'clock to 3.00 o'clock, then disappeared during a delay, and reappeared at a final stop location beyond 3.00 o'clock. After acquisition of a discrimination with final stop locations at 3.00 o'clock and 6.00 o'clock, the evidence for imagery was the accurate responding of the pigeons to novel locations at 4.00 o'clock and 7.00 o'clock. Pigeons display evidence of imagery by transforming a representation of movement that includes a series of intermediate steps which accurately represent the location of a moving stimulus after it disappears.
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Pepperberg, I. M., & Brezinsky, M. V. (1991). Acquisition of a relative class concept by an African gray parrot (Psittacus erithacus): discriminations based on relative size. J Comp Psychol, 105(3), 286–294.
Abstract: We report that an African gray parrot (Psittacus erithacus), Alex, responds to stimuli on a relative basis. Previous laboratory studies with artificial stimuli (such as pure tones) suggest that birds make relational responses as a secondary strategy, only after they have acquired information about the absolute values of the stimuli. Alex, however, after learning to respond to a small set of exemplars on the basis of relative size, transferred this behavior to novel situations that did not provide specific information about the absolute values of the stimuli. He responded to vocal questions about which was the larger or smaller exemplar by vocally labeling its color or material, and he responded “none” if the exemplars did not differ in size. His overall accuracy was 78.7%.
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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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Reid, P. J., & Shettleworth, S. J. (1992). Detection of cryptic prey: search image or search rate? J Exp Psychol Anim Behav Process, 18(3), 273–286.
Abstract: Animals' improvement in capturing cryptic prey with experience has long been attributed to a perceptual mechanism, the specific search image. Detection could also be improved by adjusting rate of search. In a series of studies using both naturalistic and operant search tasks, pigeons searched for wheat, dyed to produce 1 conspicuous and 2 equally cryptic prey types. Contrary to the predictions of the search-rate hypothesis, pigeons given a choice between the 2 cryptic types took the type experienced most recently. However, experience with 1 cryptic type improved accuracy on the other cryptic type, a result inconsistent with a search image specific to 1 prey type. Search image may better be thought of as priming of attention to those features of the prey type that best distinguish the prey from the background.
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Zentall, T. R., Roper, K. L., & Sherburne, L. M. (1995). Most directed forgetting in pigeons can be attributed to the absence of reinforcement on forget trials during training or to other procedural artifacts. J Exp Anal Behav, 63(2), 127–137.
Abstract: In research on directed forgetting in pigeons using delayed matching procedures, remember cues, presented in the delay interval between sample and comparisons, have been followed by comparisons (i.e., a memory test), whereas forget cues have been followed by one of a number of different sample-independent events. The source of directed forgetting in delayed matching to sample in pigeons was examined in a 2 x 2 design by independently manipulating whether or not forget-cue trials in training ended with reinforcement and whether or not forget-cue trials in training included a simultaneous discrimination (involving stimuli other than those used in the matching task). Results were consistent with the hypothesis that reinforced responding following forget cues is sufficient to eliminate performance deficits on forget-cue probe trials. Only when reinforcement was omitted on forget-cue trials in training (whether a discrimination was required or not) was there a decrement in accuracy on forget-cue probe trials. When reinforcement is present, however, the pattern of responding established during and following a forget cue in training may also play a role in the directed forgetting effect. These findings support the view that much of the evidence for directed forgetting using matching procedures may result from motivational and behavioral artifacts rather than the loss of memory.
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Hampton, R. R., Sherry, D. F., Shettleworth, S. J., Khurgel, M., & Ivy, G. (1995). Hippocampal volume and food-storing behavior are related in parids. Brain Behav Evol, 45(1), 54–61.
Abstract: The size of the hippocampus has been previously shown to reflect species differences and sex differences in reliance on spatial memory to locate ecologically important resources, such as food and mates. Black-capped chickadees (Parus atricapillus) cached more food than did either Mexican chickadees (P. sclateri) or bridled titmice (P. wollweberi) in two tests of food storing, one conducted in an aviary and another in smaller home cages. Black-capped chickadees were also found to have a larger hippocampus, relative to the size of the telencephalon, than the other two species. Differences in the frequency of food storing behavior among the three species have probably produced differences in the use of hippocampus-dependent memory and spatial information processing to recover stored food, resulting in graded selection for size of the hippocampus.
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Hampton, R. R., & Shettleworth, S. J. (1996). Hippocampus and memory in a food-storing and in a nonstoring bird species. Behav Neurosci, 110(5), 946–964.
Abstract: Food-storing birds maintain in memory a large and constantly changing catalog of the locations of stored food. The hippocampus of food-storing black-capped chickadees (Parus atricapillus) is proportionally larger than that of nonstoring dark-eyed juncos (Junco hyemalis). Chickadees perform better than do juncos in an operant test of spatial non-matching-to-sample (SNMTS), and chickadees are more resistant to interference in this paradigm. Hippocampal lesions attenuate performance in SNMTS and increase interference. In tests of continuous spatial alternation (CSA), juncos perform better than chickadees. CSA performance also declines following hippocampal lesions. By itself, sensitivity of a given task to hippocampal damage does not predict the direction of memory differences between storing and nonstoring species.
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