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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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Milgram, N. W., Head, E., Muggenburg, B., Holowachuk, D., Murphey, H., Estrada, J., et al. (2002). Landmark discrimination learning in the dog: effects of age, an antioxidant fortified food, and cognitive strategy. Neurosci Biobehav Rev, 26(6), 679–695.
Abstract: The landmark discrimination learning test can be used to assess the ability to utilize allocentric spatial information to locate targets. The present experiments examined the role of various factors on performance of a landmark discrimination learning task in beagle dogs. Experiments 1 and 2 looked at the effects of age and food composition. Experiments 3 and 4 were aimed at characterizing the cognitive strategies used in performance on this task and in long-term retention. Cognitively equivalent groups of old and young dogs were placed into either a test group maintained on food enriched with a broad-spectrum of antioxidants and mitochondrial cofactors, or a control group maintained on a complete and balanced food formulated for adult dogs. Following a wash-in period, the dogs were tested on a series of problems, in which reward was obtained when the animal responded selectively to the object closest to a thin wooden block, which served as a landmark. In Experiment 1, dogs were first trained to respond to a landmark placed directly on top of coaster, landmark 0 (L0). In the next phase of testing, the landmark was moved at successively greater distances (1, 4 or 10 cm) away from the reward object. Learning varied as a function of age group, food group, and task. The young dogs learned all of the tasks more quickly than the old dogs. The aged dogs on the enriched food learned L0 significantly more rapidly than aged dogs on control food. A higher proportion of dogs on the enriched food learned the task, when the distance was increased to 1cm. Experiment 2 showed that accuracy decreased with increased distance between the reward object and landmark, and this effect was greater in old animals. Experiment 3 showed stability of performance, despite using a novel landmark, and new locations, indicating that dogs learned the landmark concept. Experiment 4 found age impaired long-term retention of the landmark task. These results indicate that allocentric spatial learning is impaired in an age-dependent manner in dogs, and that age also affects performance when the distance between the landmark and target is increased. In addition, these results both support a role of oxidative damage in the development of age-associated cognitive dysfunction and indicate that short-term administration of a food enriched with supplemental antioxidants and mitochondrial cofactors can partially reverse the deleterious effects of aging on cognition.
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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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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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Wallace, D. G., Hamilton, D. A., & Whishaw, I. Q. (2006). Movement characteristics support a role for dead reckoning in organizing exploratory behavior. Anim. Cogn., 9(3), 219–228.
Abstract: Rat exploration is an organized series of trips. Each exploratory trip involves an outward tour from the refuge followed by a return to the refuge. A tour consists of a sequence of progressions with variable direction and speed concatenated by stops, whereas the return consists of a single direct progression. We have argued that processing self-movement information generated on the tour allows a rat to plot the return to the refuge. This claim has been supported by observing consistent differences between tour and return segments independent of ambient cue availability; however, this distinction was based on differences in movement characteristics derived from multiple progressions and stops on the tour and the single progression on the return. The present study examines movement characteristics of the tour and return progressions under novel-dark and light conditions. Three novel characteristics of progressions were identified: (1) linear speeds and path curvature of exploratory trips are negatively correlated, (2) tour progression maximum linear speed and temporal pacing varies as a function of travel distance, and (3) return progression movement characteristics are qualitatively different from tour progressions of comparable length. These observations support a role for dead reckoning in organizing exploratory behavior.
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Sturz, B. R., Bodily, K. D., & Katz, J. S. (2006). Evidence against integration of spatial maps in humans. Anim. Cogn., 9(3), 207–217.
Abstract: A dynamic 3-D virtual environment was constructed for humans as an open-field analogue of Blaisdell and Cook's (2005) pigeon foraging task to determine if humans, like pigeons, were capable of integrating separate spatial maps. Participants used keyboard keys and a mouse to search for a hidden goal in a 4x4 grid of raised cups. During Phase 1 training, a goal was consistently located between two landmarks (Map 1: blue T and red L). During Phase 2 training, a goal was consistently located down and left of a single landmark (Map 2: blue T). Transfer trials were then conducted in which participants were required to make choices in the presence of the red L alone. Cup choices during transfer assessed participants' strategies: association (from Map 1), generalization (from Map 2), or integration (combining Map 1 and 2). During transfer, cup choices increased to a location which suggested an integration strategy and was consistent with results obtained with pigeons. However, additional analyses of the human data suggested participants initially used a generalization strategy followed by a progressive shift in search behavior away from the red L. This shift in search behavior during transfer was responsible for the changes in cup choices across transfer trials and was confirmed by a control condition. These new analyses offer an alternative explanation to the spatial integration account proposed for pigeons.
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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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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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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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