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Bennett, A. T. (1996). Do animals have cognitive maps? J Exp Biol, 199(Pt 1), 219–224.
Abstract: Drawing on studies of humans, rodents, birds and arthropods, I show that 'cognitive maps' have been used to describe a wide variety of spatial concepts. There are, however, two main definitions. One, sensu Tolman, O'Keefe and Nadel, is that a cognitive map is a powerful memory of landmarks which allows novel short-cutting to occur. The other, sensu Gallistel, is that a cognitive map is any representation of space held by an animal. Other definitions with quite different meanings are also summarised. I argue that no animal has been conclusively shown to have a cognitive map, sensu Tolman, O'Keefe and Nadel, because simpler explanations of the crucial novel short-cutting results are invariably possible. Owing to the repeated inability of experimenters to eliminate these simpler explanations over at least 15 years, and the confusion caused by the numerous contradictory definitions of a cognitive map, I argue that the cognitive map is no longer a useful hypothesis for elucidating the spatial behaviour of animals and that use of the term should be avoided.
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Blaisdell, A. P., & Cook, R. G. (2005). Integration of spatial maps in pigeons. Anim. Cogn., 8(1), 7–16.
Abstract: The integration of spatial maps in pigeons was investigated using a spatial analog to sensory preconditioning. The pigeons were tested in an open-field arena in which they had to locate hidden food among a 4x4 grid of gravel-filled cups. In phase 1, the pigeons were exposed to a consistent spatial relationship (vector) between landmark L (a red L-shaped block of wood), landmark T (a blue T-shaped block of wood) and the hidden food goal. In phase 2, the pigeons were then exposed to landmark T with a different spatial vector to the hidden food goal. Following phase 2, pigeons were tested with trials on which they were presented with only landmark L to examine the potential integration of the phase 1 and 2 vectors via their shared common elements. When these test trials were preceded by phase 1 and phase 2 reminder trials, pigeons searched for the goal most often at a location consistent with their integration of the L-->T phase 1 and T-->phase 2 goal vectors. This result indicates that integration of spatial vectors acquired during phases 1 and 2 allowed the pigeons to compute a novel L-->goal vector. This suggests that spatial maps may be enlarged by successively integrating additional spatial information through the linkage of common elements.
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Cheng, K., & Wignall, A. E. (2006). Honeybees (Apis mellifera) holding on to memories: response competition causes retroactive interference effects. Anim. Cogn., 9(2), 141–150.
Abstract: Five experiments on honeybees examined how the learning of a second task interferes with what was previously learned. Free flying bees were tested for landmark-based memory in variations on a paradigm of retroactive interference. Bees first learned Task 1, were tested on Task 1 (Test 1), then learned Task 2, and were tested again on Task 1 (Test 2). A 60-min delay (waiting in a box) before Test 2 caused no performance decrements. If the two tasks had conflicting response requirements, (e.g., target right of a green landmark in Task 1 and left of a blue landmark in Task 2), then a strong decrement on Test 2 was found (retroactive interference effect). When response competition was minimised during training or testing, however, the decrement on Test 2 was small or nonexistent. The results implicate response competition as a major contributor to the retroactive interference effect. The honeybee seems to hold on to memories; new memories do not wipe out old ones.
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Chiesa, A. D., Pecchia, T., Tommasi, L., & Vallortigara, G. (2006). Multiple landmarks, the encoding of environmental geometry and the spatial logics of a dual brain. Anim. Cogn., 9(4), 281–293.
Abstract: A series of place learning experiments was carried out in young chicks (Gallus gallus) in order to investigate how the geometry of a landmark array and that of a walled enclosure compete when disoriented animals could rely on both of them to re-orient towards the centre of the enclosure. A square-shaped array (four wooden sticks) was placed in the middle of a square-shaped enclosure, the two structures being concentric. Chicks were trained to ground-scratch to search for food hidden in the centre of the enclosure (and the array). To check for effects of array degradation, one, two, three or all landmarks were removed during test trials. Chicks concentrated their searching activity in the central area of the enclosure, but their accuracy was inversely contingent on the number of landmarks removed; moreover, the landmarks still present within the enclosure appeared to influence the shape of the searching patterns. The reduction in the number of landmarks affected the searching strategy of chicks, suggesting that they had focussed mainly on local cues when landmarks were present within the enclosure. When all the landmarks were removed, chicks searched over a larger area, suggesting an absolute encoding of distances from the local cues and less reliance on the relationships provided by the geometry of the enclosure. Under conditions of monocular vision, chicks tended to rely on different strategies to localize the centre on the basis of the eye (and thus the hemisphere) in use, the left hemisphere attending to details of the environment and the right hemisphere attending to the global shape.
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Church, D. L., & Plowright, C. M. S. (2006). Spatial encoding by bumblebees (Bombus impatiens) of a reward within an artificial flower array. Anim. Cogn., 9(2), 131–140.
Abstract: We presented bumblebees a spatial memory task similar to that used with other species (e.g., cats, dogs, and pigeons). In some conditions we allowed for presence of scent marks in addition to placing local and global spatial cues in conflict. Bumblebees (Bombus impatiens) were presented an array of artificial flowers within a flight cage, one flower offering reward (S+), while the others were empty (S-). Bees were tested with empty flowers. In Experiment 1, flowers were either moved at the time of testing or not. Bees returned to the flower in the same absolute position of the S+ (the flower-array-independent (FAI) position), even if it was in the wrong position relative to the S- and even when new flower covers prevented the use of possible scent marks. New flower covers (i.e., without possible scent marks) had the effect of lowering the frequency of probing behavior. In Experiment 2, the colony was moved between training and testing. Again, bees chose the flower in the FAI position of the S+, and not the flower that would be chosen using strictly memory for a flight vector. Together, these experiments show that to locate the S+ bees did not rely on scent marks nor the positions of the S-, though the S- were prominent objects close to the goal. Also, bees selected environmental features to remember the position of the S+ instead of relying upon a purely egocentric point of view. Similarities with honeybees and vertebrates are discussed, as well as possible encoding mechanisms.
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Fragaszy, D., Johnson-Pynn, J., Hirsh, E., & Brakke, K. (2003). Strategic navigation of two-dimensional alley mazes: comparing capuchin monkeys and chimpanzees. Anim. Cogn., 6(3), 149–160.
Abstract: Planning is an important component of cognition that contributes, for example, to efficient movement through space. In the current study we presented novel two-dimensional alley mazes to four chimpanzees and three capuchin monkeys to identify the nature and efficiency of planning in relation to varying task parameters. All the subjects solved more mazes without error than expected by chance, providing compelling evidence that both species planned their choices in some manner. The probability of making a correct choice on mazes designed to be more demanding and presented later in the testing series was higher than on earlier, simpler mazes (chimpanzees), or unchanged (capuchin monkeys), suggesting microdevelopment of strategic choice. Structural properties of the mazes affected both species' choices. Capuchin monkeys were less likely than chimpanzees to take a correct path that initially led away from the goal but that eventually led to the goal. Chimpanzees were more likely to make an error by passing a correct path than by turning onto a wrong path. Chimpanzees and one capuchin made more errors on choices farther in sequence from the goal. Each species corrected errors before running into the end of an alley in approximately 40% of cases. Together, these findings suggest nascent planning abilities in each species, and the prospect for significant development of strategic planning capabilities on tasks presenting multiple simultaneous or sequential spatial relations. The computerized maze paradigm appears well suited to investigate movement planning and spatial perception in human and nonhuman primates alike.
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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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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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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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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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