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Hampton, R. R., & Shettleworth, S. J. (1996). Hippocampal lesions impair memory for location but not color in passerine birds. Behav Neurosci, 110(4), 831–835.
Abstract: The effects of hippocampal complex lesions on memory for location and color were assessed in black-capped chickadees (Parus atricapillus) and dark-eyed juncos (Junco hyemalis) in operant tests of matching to sample. Before surgery, most birds were more accurate on tests of memory for location than on tests of memory for color. Damage to the hippocampal complex caused a decline in memory for location, whereas memory for color was not affected in the same birds. This dissociation indicates that the avian hippocampus plays an important role in spatial cognition and suggests that this brain structure may play no role in working memory generally.
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Shettleworth, S. J. (1993). Varieties of learning and memory in animals. J Exp Psychol Anim Behav Process, 19(1), 5–14.
Abstract: It is often assumed that there is more than one kind of learning--or more than one memory system--each of which is specialized for a different function. Yet, the criteria by which the varieties of learning and memory should be distinguished are seldom clear. Learning and memory phenomena can differ from one another across species or situations (and thus be specialized) in a number of different ways. What is needed is a consistent theoretical approach to the whole range of learning phenomena, and one is explored here. Parallels and contrasts in the study of sensory systems illustrate one way to integrate the study of general mechanisms with an appreciation of species-specific adaptations.
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Reid, P. J., & Shettleworth, S. J. (1992). Detection of cryptic prey: search image or search rate? J Exp Psychol Anim Behav Process, 18(3), 273–286.
Abstract: Animals' improvement in capturing cryptic prey with experience has long been attributed to a perceptual mechanism, the specific search image. Detection could also be improved by adjusting rate of search. In a series of studies using both naturalistic and operant search tasks, pigeons searched for wheat, dyed to produce 1 conspicuous and 2 equally cryptic prey types. Contrary to the predictions of the search-rate hypothesis, pigeons given a choice between the 2 cryptic types took the type experienced most recently. However, experience with 1 cryptic type improved accuracy on the other cryptic type, a result inconsistent with a search image specific to 1 prey type. Search image may better be thought of as priming of attention to those features of the prey type that best distinguish the prey from the background.
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Shettleworth, S. J., & Plowright, C. M. (1992). How pigeons estimate rates of prey encounter. J Exp Psychol Anim Behav Process, 18(3), 219–235.
Abstract: Pigeons were trained on operant schedules simulating successive encounters with prey items. When items were encountered on variable-interval schedules, birds were more likely to accept a poor item (long delay to food) the longer they had just searched, as if they were averaging prey density over a short memory window (Experiment 1). Responding as if the immediate future would be like the immediate past was reversed when a short search predicted a long search next time (Experiment 2). Experience with different degrees of environmental predictability appeared to change the length of the memory window (Experiment 3). The results may reflect linear waiting (Higa, Wynne, & Staddon, 1991), but they differ in some respects. The findings have implications for possible mechanisms of adjusting behavior to current reinforcement conditions.
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Shettleworth, S. J. (1985). Foraging, memory, and constraints on learning. Ann N Y Acad Sci, 443, 216–226.
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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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Shettleworth, S. J. (1972). Stimulus relevance in the control of drinking and conditioned fear responses in domestic chicks (Gallus gallus). J Comp Physiol Psychol, 80(2), 175–198.
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Nevin, J. A., & Shettleworth, S. J. (1966). An analysis of contrast effects in multiple schedules. J Exp Anal Behav, 9(4), 305–315.
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Mendl, M. (1999). Performing under pressure: stress and cognitive function. Appl. Anim. Behav. Sci., 65(3), 221–244.
Abstract: The way in which cognitive functioning is affected by stressors is an important area of research for applied ethologists because stress caused by captive conditions may disrupt cognitive processes and lead to welfare and husbandry problems. Such problems may be minimised through an understanding of the links between stress and cognition. The effects of stress on cognitive function have been studied in disciplines ranging from human perceptual psychology to animal neuroscience. The aim of this paper is to provide an introduction to this research, focusing on the effects of stressors on attention, memory formation and memory recall. Findings from such a diverse literature with little apparent inter-disciplinary communication are inevitably complex and often contradictory. Nevertheless, some generalities do emerge. The idea that an inverted U-shaped relationship exists between an individual's state of stress or arousal and its ability to perform a cognitive task effectively, the so-called Yerkes-Dodson law, is commonly encountered. The law has limited explanatory value because it is unlikely that different stressors act on cognitive function via the same intervening, non-specific state. Furthermore, the law only provides a very general description of the relationship between stress and cognitive function. Empirical research on attention and memory processes reveals more specific findings. Stressors appear to cause shifts, lapses and narrowing of attention, and can also influence decision speed. These processes may be viewed as serving an adaptive role helping the animal to search for and scrutinise a source of danger. There is conflicting evidence as to whether hormones involved in the hypothalamic-pituitary-adrenal stress response play a part in these processes. These hormones and those involved in the sympathetic-adrenomedullary stress response do appear to play an important role in memory formation. Low or moderate concentrations of circulating glucocorticoids and catecholamines can enhance memory formation, while excessively high or prolonged elevations of these hormones can lead to memory disruption. The effects of stressors on memory recall are less clear. There is evidence for disruptive effects, and for facilitatory effects indicating state-dependent memory recall; events experienced under conditions of high arousal may be best recalled under similar conditions. Applied ethologists have the opportunity to extend work in this area, which often involves studies of single stressors/stress hormones acting in isolation and limited measures of cognitive function, by focusing on real-life husbandry stressors encountered by captive animals. This will yield fundamental information which also has direct relevance to animal welfare and management issues.
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Hanggi, E. B. (2003). Discrimination learning based on relative size concepts in horses (Equus caballus). Appl. Anim. Behav. Sci., 83(3), 201–213.
Abstract: This study explored whether or not horses (Equus caballus) could respond to stimuli using a concept based on relative size. In Experiment 1, after learning to respond to the larger of the two stimuli for six sets of two-dimensional (2D) training exemplars, one horse was tested for size transposition that used novel larger and smaller stimuli as well as three-dimensional (3D) objects (5 two-dimensional sets and 5 three-dimensional sets with large, medium, small, and tiny sizes). The horse correctly chose (significantly above chance) the larger of two stimuli regardless of novelty or dimension or combination. In Experiment 2, two additional horses were tested using a subset of the stimuli from Experiment 1. One horse was required to select the larger stimulus--as in Experiment 1--and the other the smaller stimulus. After learning the task, both horses responded correctly to new stimuli and showed size transposition. These results suggest that at least some horses are capable of solving problems based on relative size concepts. Moreover, they are able to generalize across situations that vary from flat, black shapes to objects of different materials and colors including balls, flower pots, and PVC connectors. These findings support earlier research that showed that horses could categorize certain stimuli, and provide new evidence that they are capable of using some form of concept for problem solving. Understanding that horses have more advanced learning abilities than was previously believed should help improve training methods and management.
Keywords: Horse; Concept; Size transposition; Generalization; Learning; Training
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