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Shettleworth, S. J. (1993). Varieties of learning and memory in animals. J Exp Psychol Anim Behav Process, 19(1), 5–14.
Abstract: It is often assumed that there is more than one kind of learning--or more than one memory system--each of which is specialized for a different function. Yet, the criteria by which the varieties of learning and memory should be distinguished are seldom clear. Learning and memory phenomena can differ from one another across species or situations (and thus be specialized) in a number of different ways. What is needed is a consistent theoretical approach to the whole range of learning phenomena, and one is explored here. Parallels and contrasts in the study of sensory systems illustrate one way to integrate the study of general mechanisms with an appreciation of species-specific adaptations.
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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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Gibson, B. M., & Shettleworth, S. J. (2005). Place versus response learning revisited: tests of blocking on the radial maze. Behav Neurosci, 119(2), 567–586.
Abstract: Neurobiological and behavioral research indicates that place learning and response learning occur simultaneously, in parallel. Such findings seem to conflict with theories of associative learning in which different cues compete for learning. The authors conducted place+response training on a radial maze and then tested place learning and response learning separately by reconfiguring the maze in various ways. Consistent with the effects of manipulating place and response systems in the brain (M. G. Packard & J. L. McGaugh, 1996), well-trained rats showed strong place learning and strong response learning. Three experiments using associative blocking paradigms indicated that prior response learning interferes with place learning. Blocking and related tests can be used to better understand how memory systems interact during learning.
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Sekuler, A. B., Lee, J. A., & Shettleworth, S. J. (1996). Pigeons do not complete partly occluded figures. Perception, 25(9), 1109–1120.
Abstract: One of the most common obstacles to object perception is the fact that objects often occlude parts of themselves and parts of other objects. Perceptual completion has been studied extensively in humans, and researchers have shown that humans do complete partly occluded objects. In an effort to understand more about the mechanisms underlying completion, recent research has extended the study of perceptual completion to other mammalian species. Monkeys and mice also seem to complete two-dimensional representations of partly occluded objects. The present study addresses the question of whether this capacity generalizes to a nonmammalian species, the pigeon (Columba livia). The results point to a limit of the generalizability of perceptual completion: pigeons do not complete partly occluded figures.
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Sole, L. M., Shettleworth, S. J., & Bennett, P. J. (2003). Uncertainty in pigeons. Psychon Bull Rev, 10(3), 738–745.
Abstract: Pigeons classified a display of illuminated pixels on a touchscreen as sparse or dense. Correct responses were reinforced with six food pellets; incorrect responses were unreinforced. On some trials an uncertain response option was available. Pecking it was always reinforced with an intermediate number of pellets. Like monkeys and people in related experiments, the birds chose the uncertain response most often when the stimulus presented was difficult to classify correctly, but in other respects their behavior was not functionally similar to human behavior based on conscious uncertainty or to the behavior of monkeys in comparable experiments. Our data were well described by a signal detection model that assumed that the birds were maximizing perceived reward in a consistent way across all the experimental conditions.
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Shettleworth, S. J., & Plowright, C. M. (1992). How pigeons estimate rates of prey encounter. J Exp Psychol Anim Behav Process, 18(3), 219–235.
Abstract: Pigeons were trained on operant schedules simulating successive encounters with prey items. When items were encountered on variable-interval schedules, birds were more likely to accept a poor item (long delay to food) the longer they had just searched, as if they were averaging prey density over a short memory window (Experiment 1). Responding as if the immediate future would be like the immediate past was reversed when a short search predicted a long search next time (Experiment 2). Experience with different degrees of environmental predictability appeared to change the length of the memory window (Experiment 3). The results may reflect linear waiting (Higa, Wynne, & Staddon, 1991), but they differ in some respects. The findings have implications for possible mechanisms of adjusting behavior to current reinforcement conditions.
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Gibson, B. M., & Shettleworth, S. J. (2003). Competition among spatial cues in a naturalistic food-carrying task. Learn Behav, 31(2), 143–159.
Abstract: Rats collected nuts from a container in a large arena in four experiments testing how learning about a beacon or cue at a goal interacts with learning about other spatial cues (place learning). Place learning was quick, with little evidence of competition from the beacon (Experiments 1 and 2). Rats trained to approach a beacon regardless of its location were subsequently impaired when the well-learned beacon was removed and other spatial cues identified the location of the goal (Experiment 3). The competition between beacon and place cues reflected learned irrelevance for place cues (Experiment 4). The findings differ from those of some studies of associative interactions between cue and place learning in other paradigms.
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Shettleworth, S. J., & Sutton, J. E. (2005). Multiple systems for spatial learning: dead reckoning and beacon homing in rats. J Exp Psychol Anim Behav Process, 31(2), 125–141.
Abstract: Rats homed with food in a large lighted arena. Without visual cues, they used dead reckoning. When a beacon indicated the home, rats could also use the beacon. Homing did not differ in 2 groups of rats, 1 provided with the beacon and 1 without it; tests without the beacon gave no evidence that beacon learning overshadowed dead reckoning (Experiment 1). When the beacon was at the home for 1 group and in random locations for another, there was again no evidence of cue competition (Experiment 2). Dead reckoning experience did not block acquisition of beacon homing (Experiment 3). Beacon learning and dead reckoning do not compete for predictive value but acquire information in parallel and are used hierarchically.
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Skov-Rackette, S. I., & Shettleworth, S. J. (2005). What do rats learn about the geometry of object arrays? Tests with exploratory behavior. J Exp Psychol Anim Behav Process, 31(2), 142–154.
Abstract: Six experiments using habituation of exploratory behavior tested whether disoriented rats foraging in a large arena encode the shapes of arrays of objects. Rats did not respond to changes in position of a single object, but they responded to a change in object color and to a change in position of 1 object in a square array, as in previous research (e.g., C. Thinus-Blanc et al., 1987). Rats also responded to an expansion of a square array, suggesting that they encoded sets of interobject distances rather than overall shape. In Experiments 4-6, rats did not respond to changes in sense of a triangular array that maintained interobject distances and angles. Shapes of object arrays are encoded differently from shapes of enclosures.
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Skov-Rackette, S. I., Miller, N. Y., & Shettleworth, S. J. (2006). What-where-when memory in pigeons. J Exp Psychol Anim Behav Process, 32(4), 345–358.
Abstract: The authors report a novel approach to testing episodic-like memory for single events. Pigeons were trained in separate sessions to match the identity of a sample on a touch screen, to match its location, and to report on the length of the retention interval. When these 3 tasks were mixed randomly within sessions, birds were more than 80% correct on each task. However, performance on 2 different tests in succession after each sample was not consistent with an integrated memory for sample location, time, and identity. Experiment 2 tested binding of location and identity memories in 2 different ways. The results were again consistent with independent feature memories. Implications for tests of episodic-like memory are discussed.
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