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Hamilton, W. D. (1971). Geometry for the selfish herd. J. Theor. Biol., 31(2), 295–311.
Abstract: This paper presents an antithesis to the view that gregarious behaviour is evolved through benefits to the population or species. Following Galton (1871) and Williams (1964) gregarious behaviour is considered as a form of cover-seeking in which each animal tries to reduce its chance of being caught by a predator.
It is easy to see how pruning of marginal individuals can maintain centripetal instincts in already gregarious species; some evidence that marginal pruning actually occurs is summarized. Besides this, simply defined models are used to show that even in non-gregarious species selection is likely to favour individuals who stay close to others.
Although not universal or unipotent, cover-seeking is a widespread and important element in animal aggregation, as the literature shows. Neglect of the idea has probably followed from a general disbelief that evolution can be dysgenic for a species. Nevertheless, selection theory provides no support for such disbelief in the case of species with outbreeding or unsubdivided populations.
The model for two dimensions involves a complex problem in geometrical probability which has relevance also in metallurgy and communication science. Some empirical data on this, gathered from random number plots, is presented as of possible heuristic value.
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Vlasak, A. N. (2006). Global and local spatial landmarks: their role during foraging by Columbian ground squirrels (Spermophilus columbianus). Anim. Cogn., 9(1), 71–80.
Abstract: Locating food and refuge is essential for an animal's survival. However, little is known how mammals navigate under natural conditions and cope with given environmental constraints. In a series of six experiments, I investigated landmark-based navigation in free-ranging Columbian ground squirrels (Spermophilus columbianus). Squirrels were trained individually to find a baited platform within an array of nine identical platforms and artificial landmarks set up on their territories. After animals learned the location of the food platform in the array, the position of the latter with respect to local artificial, local natural, and global landmarks was manipulated, and the animal's ability to find the food platform was tested. When only positions of local artificial landmarks were changed, squirrels located food with high accuracy. When the location of the array relative to global landmarks was altered, food-finding accuracy decreased but remained significant. In the absence of known global landmarks, the presence of a familiar route and natural local landmarks resulted in significant but not highly accurate performance. Squirrels likely relied on multiple types of cues when orienting towards a food platform. Local landmarks were used only as a secondary mechanism of navigation, and were not attended to when a familiar route and known global landmarks were present. This study provided insights into landmark use by a wild mammal in a natural situation, and it demonstrated that an array of platforms can be employed to investigate landmark-based navigation under such conditions.
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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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Range, F., Bugnyar, T., Schlogl, C., & Kotrschal, K. (2006). Individual and sex differences in learning abilities of ravens. Behav. Process., 73(1), 100–106.
Abstract: Behavioral and physiological characteristics of individuals within the same species have been found to be stable across time and contexts. In this study, we investigated individual differences in learning abilities and object and social manipulation to test for consistency within individuals across different tasks. Individual ravens (Corvus corax) were tested in simple color and position discrimination tasks to establish their learning abilities. We found that males were significantly better in the acquisition of the first discrimination task and the object manipulation task, but not in any of the other tasks. Furthermore, faster learners engaged less often in manipulations of conspecifics and exploration of objects to get access to food. No relationship between object and social manipulation and reversal training were found. Our results suggest that individual differences in regard to the acquisition of new tasks may be related to personalities or at least object manipulation in ravens.
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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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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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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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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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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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Jensen, G. D., Gordon, B. N., & Wolfheim, J. (1975). Nursing behavior in infant monkeys: a sequence analysis. Behaviour, 55(1-2), 115–127.
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