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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Hampton, R. R., Healy, S. D., Shettleworth, S. J., & Kamil, A. C. (2002). Neuroecologists' are not made of straw. Trends. Cognit. Sci., 6(1), 6–7.
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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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Shettleworth, S. J. (2003). Memory and hippocampal specialization in food-storing birds: challenges for research on comparative cognition. Brain Behav Evol, 62(2), 108–116.
Abstract: The three-way association among food-storing behavior, spatial memory, and hippocampal enlargement in some species of birds is widely cited as an example of a new 'cognitive ecology' or 'neuroecology.' Whether this relationship is as strong as it first appears and whether it might be evidence for an adaptive specialization of memory and hippocampus in food-storers have recently been the subject of some controversy [Bolhuis and Macphail, 2001; Macphail and Bolhuis, 2001]. These critiques are based on misconceptions about the nature of adaptive specializations in cognition, misconceptions about the uniformity of results to be expected from applying the comparative method to data from a wide range of species, and a narrow view of what kinds of cognitive adaptations are theoretically interesting. New analyses of why food-storers (black-capped chickadees, Poecile Atricapilla) respond preferentially to spatial over color cues when both are relevant in a memory task show that this reflects a relative superiority of spatial memory as compared to memory for color rather than exceptional spatial attention or spatial discrimination ability. New studies of chickadees from more or less harsh winter climates also support the adaptive specialization hypothesis and suggest that within-species comparisons may be especially valuable for unraveling details of the relationships among ecology, memory, and brain in food-storing species.
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Gibson, B. M., Juricevic, I., Shettleworth, S. J., Pratt, J., & Klein, R. M. (2005). Looking for inhibition of return in pigeons. Learn Behav, 33(3), 296–308.
Abstract: We conducted four experiments in order to investigate whether pigeons' responses to a recently attended (i.e., recently pecked) location are inhibited. In Experiments 1 and 2, stimulus displays were similar to those used in studies of inhibition of return (IOR) with humans; responses to cued targets tended to be facilitated rather than inhibited. In Experiments 3 and 4, birds were presented with stimulus displays that mimicked clusters of small grains and were relatively localized, which should have been more appropriate for detecting IOR in pigeons. The results from these experiments again provided evidence for facilitation of responding to cued targets, rather than for IOR.
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Sutton, J. E., & Shettleworth, S. J. (2005). Internal sense of direction and landmark use in pigeons (Columba livia). J Comp Psychol, 119(3), 273–284.
Abstract: The relative importance of an internal sense of direction based on inertial cues and landmark piloting for small-scale navigation by White King pigeons (Columba livia) was investigated in an arena search task. Two groups of pigeons differed in whether they had access to visual cues outside the arena. In Experiment 1, pigeons were given experience with 2 different entrances and all pigeons transferred accurate searching to novel entrances. Explicit disorientation before entering did not affect accuracy. In Experiments 2-4, landmarks and inertial cues were put in conflict or tested 1 at a time. Pigeons tended to follow the landmarks in a conflict situation but could use an internal sense of direction to search when landmarks were unavailable.
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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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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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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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