Shettleworth, S. J. (1978). Reinforcement and the organization of behavior in golden hamsters: Pavlovian conditioning with food and shock unconditioned stimuli. J Exp Psychol Anim Behav Process, 4(2), 152–169.
Abstract: The effects of Pavlovian conditioned stimuli (CSs) for food or shock on a variety of behaviors of golden hamsters were observed in three experiments. The aim was to see whether previously reported differences among the behaviors produced by food reinforcement and punishment procedures could be accounted for by differential effects of Pavlovian conditioning on the behaviors. There was some correspondence between the behaviors observed to the CSs and the previously reported effects of instrumental training. However, the Pavlovian conditioned responses (CRs) alone would not have predicted the effects of instrumental training. Moreover, CRs depended to some extent on the context in which training and testing occurred. These findings, together with others in the literature, suggest that the results of Pavlovian conditioning procedures may not unambiguously predict what system of behaviors will be most readily modified by instrumental training with a given reinforcer.
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Shettleworth, S. J., & Juergensen, M. R. (1980). Reinforcement and the organization of behavior in golden hamsters: brain stimulation reinforcement for seven action patterns. J Exp Psychol Anim Behav Process, 6(4), 352–375.
Abstract: Golden hamsters were reinforced with intracranial electrical stimulation of the lateral hypothalamus (ICS) for spending time engaging in one of seven topographically defined action patterns (APs). The stimulation used as reinforcer elicited hoarding and/or feeding and supported high rates of bar pressing. In Experiment 1, hamsters were reinforced successively for digging, open rearing, and face washing. Digging increased most in time spent, and face washing increased least. Experiments 2-5 examined these effects further and also showed that “scrabbling,” like digging, was performed a large proportion of the time, almost without interruption, for contingent ICS but that scratching the body with a hindleg and scent-marking showed relatively little effect of contingent ICS, the latter even in an environment that facilitated marking. In Experiment 6, naive hamsters received ICS not contingent on behavior every 30 sec (fixed-time 30-sec schedule). Terminal behaviors that developed on this schedule were APs that were easy to reinforce in the other experiments, but a facultative behavior, face washing, was one not so readily reinforced. Experiment 7 confirmed a novel prediction from Experiment 6--that wall rearing, a terminal AP, would be performed at a high level for contingent ICS. All together, the results point to both motivational factors and associative factors being involved in the considerable differences in performance among different reinforced activities.
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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., 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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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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Krebs, J. R., Clayton, N. S., Hampton, R. R., & Shettleworth, S. J. (1995). Effects of photoperiod on food-storing and the hippocampus in birds. Neuroreport, 6(12), 1701–1704.
Abstract: Birds that store food have a relatively large hippocampus compared to non-storing species. The hippocampus shows seasonal differences in neurogenesis and volume in black-capped chikadees (Parus atricapillus) taken from the wild at different times of year. We compared hippocampal volumes in black-capped chickadees captured at the same time but differing in food-storing behaviour because of manipulations of photoperiod in the laboratory. Differences in food-storing behaviour were not accompanied by differences in the volume of the hippocampus. Hippocampal volumes also did not differ between two groups of a non-food-storing control species, house sparrows (Passer domesticus), exposed to the same conditions as the chickadees.
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Hampton, R. R., & Shettleworth, S. J. (1996). Hippocampal lesions impair memory for location but not color in passerine birds. Behav Neurosci, 110(4), 831–835.
Abstract: The effects of hippocampal complex lesions on memory for location and color were assessed in black-capped chickadees (Parus atricapillus) and dark-eyed juncos (Junco hyemalis) in operant tests of matching to sample. Before surgery, most birds were more accurate on tests of memory for location than on tests of memory for color. Damage to the hippocampal complex caused a decline in memory for location, whereas memory for color was not affected in the same birds. This dissociation indicates that the avian hippocampus plays an important role in spatial cognition and suggests that this brain structure may play no role in working memory generally.
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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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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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