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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., & 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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Petruso, E. J., Fuchs, T., & Bingman, V. P. (2007). Time-space learning in homing pigeons (Columba livia): orientation to an artificial light source. Anim. Cogn., 10(2), 181–188.
Abstract: Time-space learning reflects an ability to represent in memory event-stimulus properties together with the place and time of the event; a capacity well developed in birds. Homing pigeons were trained in an indoor octagonal arena to locate one food goal in the morning and a different food goal in the late afternoon. The goals differed with respect to their angular/directional relationship to an artificial light source located outside the arena. Further, the angular difference in reward position approximated the displacement of the sun's azimuth that would occur during the same time period. The experimental birds quickly learned the task, demonstrating the apparent ease with which birds can adopt an artificial light source to discriminate among alternative spatial responses at different times of the day. However, a novel midday probe session following successful learning revealed that the light source was interpreted as a stable landmark and not as a surrogate sun that would support compass orientation. Probe sessions following a phase shift of the light-dark cycle revealed that the mechanism employed to make the temporal discrimination was prevailingly based on an endogenous circadian rhythm and not an interval timing mechanism.
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Merchant, H., Fortes, A. F., & Georgopoulos, A. P. (2004). Short-term memory effects on the representation of two-dimensional space in the rhesus monkey. Anim. Cogn., 7(3), 133–143.
Abstract: Human subjects represent the location of a point in 2D space using two independent dimensions (x-y in Euclidean or radius-angle in polar space), and encode location in memory along these dimensions using two levels of representation: a fine-grain value and a category. Here we determined whether monkeys possessed the ability to represent location with these two levels of coding. A rhesus monkey was trained to reproduce the location of a dot in a circle by pointing, after a delay period, on the location where a dot was presented. Five different delay periods (0.5-5 s) were used. The results showed that the monkey used a polar coordinate system to represent the fine-grain spatial coding, where the radius and angle of the dots were encoded independently. The variability of the spatial response and reaction time increased with longer delays. Furthermore, the animal was able to form a categorical representation of space that was delay-dependent. The responses avoided the circumference and the center of the circle, defining a categorical radial prototype around one third of the total radial length. This radial category was observed only at delay durations of 3-5 s. Finally, the monkey also formed angular categories with prototypes at the obliques of the quadrants of the circle, avoiding the horizontal and vertical axes. However, these prototypes were only observed at the 5-s delay and on dots lying on the circumference. These results indicate that monkeys may possess spatial cognitive abilities similar to humans.
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Kelly, D. M., & Spetch, M. L. (2001). Pigeons encode relative geometry. J Exp Psychol Anim Behav Process, 27(4), 417–422.
Abstract: Pigeons were trained to search for hidden food in a rectangular environment designed to eliminate any external cues. Following training, the authors administered unreinforced test trials in which the geometric properties of the apparatus were manipulated. During tests that preserved the relative geometry but altered the absolute geometry of the environment, the pigeons continued to choose the geometrically correct corners, indicating that they encoded the relative geometry of the enclosure. When tested in a square enclosure, which distorted both the absolute and relative geometry, the pigeons randomly chose among the 4 corners, indicating that their choices were not based on cues external to the apparatus. This study provides new insight into how metric properties of an environment are encoded by pigeons.
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Hodgson, Z. G., & Healy, S. D. (2005). Preference for spatial cues in a non-storing songbird species. Anim. Cogn., 8(3), 211–214.
Abstract: Male mammals typically outperform their conspecific females on spatial tasks. A sex difference in cues used to solve the task could underlie this performance difference as spatial ability is reliant on appropriate cue use. Although comparative studies of memory in food-storing and non-storing birds have examined species differences in cue preference, few studies have investigated differences in cue use within a species. In this study, we used a one-trial associative food-finding task to test for sex differences in cue use in the great tit, Parus major. Birds were trained to locate a food reward hidden in a well covered by a coloured cloth. To determine whether the colour of the cloth or the location of the well was learned during training, the birds were presented with three wells in the test phase: one in the original location, but covered by a cloth of a novel colour, a second in a new location covered with the original cloth and a third in a new location covered by a differently coloured cloth. Both sexes preferentially visited the well in the training location rather than either alternative. As great tits prefer colour cues over spatial cues in one-trial associative conditioning tasks, cue preference appears to be related to the task type rather than being species dependent.
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Herrmann, E., Melis, A. P., & Tomasello, M. (2006). Apes' use of iconic cues in the object-choice task. Anim. Cogn., 9(2), 118–130.
Abstract: In previous studies great apes have shown little ability to locate hidden food using a physical marker placed by a human directly on the target location. In this study, we hypothesized that the perceptual similarity between an iconic cue and the hidden reward (baited container) would help apes to infer the location of the food. In the first two experiments, we found that if an iconic cue is given in addition to a spatial/indexical cue – e.g., picture or replica of a banana placed on the target location – apes (chimpanzees, bonobos, orangutans, gorillas) as a group performed above chance. However, we also found in two further experiments that when iconic cues were given on their own without spatial/indexical information (iconic cue held up by human with no diagnostic spatial/indexical information), the apes were back to chance performance. Our overall conclusion is that although iconic information helps apes in the process of searching hidden food, the poor performance found in the last two experiments is due to apes' lack of understanding of the informative (cooperative) communicative intention of the experimenter.
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Henderson, J., Hurly, T. A., & Healy, S. D. (2006). Spatial relational learning in rufous hummingbirds (Selasphorus rufus). Anim. Cogn., 9(3), 201–205.
Abstract: There is increasing evidence that animals can learn abstract spatial relationships, and successfully transfer this knowledge to novel situations. In this study, rufous hummingbirds (Selasphorus rufus) were trained to feed from either the lower or the higher of two flowers. When presented with a test pair of flowers, one of which was at a novel height, they chose the flower in the appropriate spatial position rather than the flower at the correct height. This response may also have been influenced by a preference for taller flowers as acquisition of the task during experimental training occurred more readily when the reward flower was the taller of the pair. Thus, it appears that although learning abstract relationships may be a general phenomenon across contexts, and perhaps across species, the ease with which they are learned and the context in which they are subsequently used may not be the same.
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Held, S., Baumgartner, J., Kilbride, A., Byrne, R. W., & Mendl, M. (2005). Foraging behaviour in domestic pigs (Sus scrofa): remembering and prioritizing food sites of different value. Anim. Cogn., 8(2), 114–121.
Abstract: This experiment investigated whether domestic pigs can remember the locations of food sites of different relative value, and how a restricted retrieval choice affects their foraging behaviour. Nine juvenile female pigs were trained to relocate two food sites out of a possible eight in a spatial memory task. The two baited sites contained different amounts of food and an obstacle was added to the smaller amount to increase handling time. On each trial, a pig searched for the two baited sites (search visit). Once it had found and eaten the bait, it returned for a second (relocation) visit, in which the two same sites were baited. Baited sites were changed between trials. All subjects learnt the task. When allowed to retrieve both baits, the subjects showed no preference for retrieving a particular one first (experiment 1). When they were allowed to retrieve only one bait, a significant overall preference for retrieving the larger amount emerged across subjects (experiment 2). To test whether this preference reflected an avoidance of the obstacle with the smaller bait, 15 choice-restricted control trials were conducted. In control trials obstacles were present with both baits. Pigs continued to retrieve the larger bait, indicating they had discriminated between the two food sites on the basis of quantity or profitability and adjusted their behaviour accordingly when the relocation choice was restricted. This suggests for the first time that domestic pigs have the ability to discriminate between food sites of different relative value and to remember their respective locations.
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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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