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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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., & 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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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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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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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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Gibbs, S. E. B., Lea, S. E. G., & Jacobs, L. F. (2007). Flexible use of spatial cues in the southern flying squirrel (Glaucomys volans). Anim. Cogn., 10(2), 203–209.
Abstract: Insects, birds, and mammals have been shown capable of encoding spatial information in memory using multiple strategies or frames of reference simultaneously. These strategies include orientation to a goal-specific cue or beacon, to the position of the goal in an array of local landmarks, or to its position in the array of distant landmarks, also known as the global frame of reference. From previous experiments, it appears that birds and mammals that scatter hoard rely primarily on a global frame of reference, but this generalization depends on evidence from only a few species. Here we examined spatial memory in a previously unstudied scatter hoarder, the southern flying squirrel. We dissociated the relative weighting of three potential spatial strategies (beacon, global, or relative array strategy) with three probe tests: transposition of beacon and the rotation or the expansion of the array. The squirrels' choices were consistent with a spatial averaging strategy, where they chose the location dictated by at least two of the three strategies, rather than using a single preferred frame of reference. This adaptive and flexible heuristic has not been previously described in animal orientation studies, yet it may be a common solution to the universal problem of encoding and recalling spatial locations in an ephemeral physical landscape.
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Fiset, S., & Leblanc, V. (2007). Invisible displacement understanding in domestic dogs (Canis familiaris): the role of visual cues in search behavior. Anim. Cogn., 10(2), 211–224.
Abstract: Recently, (Collier-Baker E, Davis JM, Suddendorf T (2004) J Comp Psychol 118:421-433) suggested that domestic dogs do not understand invisible displacements. In the present study, we further investigated the hypothesis that the search behavior of domestic dogs in invisible displacements is guided by various visual cues inherent to the task rather than by mental representation of an object's past trajectory. Specifically, we examined the role of the experimenter as a function of the final position of the displacement device in the search behavior of domestic dogs. Visible and invisible displacement problems were administered to dogs (N = 11) under two conditions. In the Visible-experimenter condition, the experimenter was visible whereas in the Concealed-experimenter condition, the experimenter was visibly occluded behind a large rigid barrier. Our data supported the conclusion that dogs do not understand invisible displacements but primarily search as a function of the final position of the displacement device and, to a lesser extent, the position of the experimenter.
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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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Chiandetti, C., Regolin, L., Sovrano, V. A., & Vallortigara, G. (2007). Spatial reorientation: the effects of space size on the encoding of landmark and geometry information. Anim. Cogn., 10(2), 159–168.
Abstract: The effects of the size of the environment on animals' spatial reorientation was investigated. Domestic chicks were trained to find food in a corner of either a small or a large rectangular enclosure. A distinctive panel was located at each of the four corners of the enclosures. After removal of the panels, chicks tested in the small enclosure showed better retention of geometrical information than chicks tested in the large enclosure. In contrast, after changing the enclosure from a rectangular-shaped to a square-shaped one, chicks tested in the large enclosure showed better retention of landmark (panels) information than chicks tested in the small enclosure. No differences in the encoding of the overall arrangement of landmarks were apparent when chicks were tested for generalisation in an enclosure differing from that of training in size together with a transformation (affine transformation) that altered the geometric relations between the target and the shape of the environment. These findings suggest that primacy of geometric or landmark information in reorientation tasks depends on the size of the experimental space, likely reflecting a preferential use of the most reliable source of information available during visual exploration of the environment.
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