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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. (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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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., & 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., & Westwood, R. P. (2002). Divided attention, memory, and spatial discrimination in food-storing and nonstoring birds, black-capped chickadees (Poecile atricapilla) and dark-eyed juncos (Junco hyemalis). J Exp Psychol Anim Behav Process, 28(3), 227–241.
Abstract: Food-storing birds, black-capped chickadees (Poecile atricapilla), and nonstoring birds, dark-eyed juncos (Junco hyemalis), matched color or location on a touch screen. Both species showed a divided attention effect for color but not for location (Experiment 1). Chickadees performed better on location than on color with retention intervals up to 40 s, but juncos did not (Experiment 2). Increasing sample-distractor distance improved performance similarly in both species. Multidimensional scaling revealed that both use a Euclidean metric of spatial similarity (Experiment 3). When choosing between the location and color of a remembered item, food storers choose location more than do nonstorers. These results explain this effect by differences in memory for location relative to color, not division of attention or spatial discrimination ability.
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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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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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Tommasi, L., & Vallortigara, G. (2000). Searching for the center: spatial cognition in the domestic chick (Gallus gallus). J Exp Psychol Anim Behav Process, 26(4), 477–486.
Abstract: Chicks learned to find food hidden under sawdust by ground-scratching in the central position of the floor of a closed arena. When tested inan arena of identical shape but a larger area, chicks searched at 2 different locations, one corresponding to the correct distance (i.e., center) in the smaller (training) arena and the other to the actual center of the test arena. When tested in an arena of the same shape but a smaller area, chicks searched in the center of it. These results suggest that chicks are able to encode information on the absolute and relative distance of the food from the walls of the arena. After training in the presence of a landmark located at the center of the arena, animals searched at the center even after the removal of the landmark. Marked changes in the height of the walls of the arena produced some displacement in searching behavior, suggesting that chicks used the angular size of the walls to estimate distances.
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