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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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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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Cooper, J. J., Ashton, C., Bishop, S., West, R., Mills, D. S., & Young, R. J. (2003). Clever hounds: social cognition in the domestic dog (Canis familiaris). Appl. Anim. Behav. Sci., 81(3), 229–244.
Abstract: This paper reviews the reasons why domestic dogs make good models to investigate cognitive processes related to social living and describes experimental approaches that can be adopted to investigate such processes in dogs. Domestic dogs are suitable models for investigating social cognition skills for three broad reasons. First, dogs originated from wolves, social animals that engage in a number of co-operative behaviours, such as hunting and that may have evolved cognitive abilities that help them predict and interpret the actions of other animals. Second, during domestication dogs are likely to have been selected for mental adaptations for their roles in human society such as herding or companionship. Third, domestic dogs live in a human world and “enculturation” may facilitate the development of relevant mental skills in dogs. Studies of social cognition in animals commonly use experimental paradigms originally developed for pre-verbal human infants. Preferential gaze, for example, can be used as a measure of attention or “surprise” in studies using expectancy violation. This approach has been used to demonstrate simple numerical competence in dogs. Dogs also readily use both conspecific and human social signals (e.g. looking or pointing) as information sources to locate hidden rewards such as food or favourite toys. Such abilities make dogs particularly good models for investigating perspective-taking tasks, where animals are required to discriminate between apparently knowledgeable and apparently ignorant informants.
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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.
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Saslow, C. A. (2002). Understanding the perceptual world of horses. Appl. Anim. Behav. Sci., 78(2-4), 209–224.
Abstract: From the viewpoint of experimental psychology, there are two problems with our current knowledge of equine perception. The first is that the behavioral and neurophysiological research in this area has enormous gaps, reflecting that this animal is not a convenient laboratory subject. The second is that the horse, having been a close companion to humans for many millennia, entrenched anecdotal wisdom is often hard to separate from scientific fact. Therefore, any summary at present of equine perception has to be provisional. The horse appears to have developed a visual system particularly sensitive to dim light and movement, it may or may not have a weak form of color vision in part of the retina, it has little binocular overlap, and its best acuity is limited to a restricted horizontal band which is aimed primarily by head/neck movements. However, the total field of view is very large. Overall, as would be expected for a prey animal, horse vision appears to have evolved more for detection of predator approach from any angle than for accurate visual identification of stationary objects, especially those seen at a distance. It is likely that, as for most mammals except the primates, horses rely more heavily on their other senses for forming a view of their world. Equine high-frequency hearing extends far above that of humans, but horses may be less able to localize the point of origin of brief sounds. The horse's capacity for chemoreception and its reliance on chemical information for identification may more closely resemble that of the dog than of the human. Its tactile sensitivity is high, and the ability of its brain and body to regulate pain perception appears to be similar to that found in other mammals. There is room for a great deal of future research in both the area of equine perception and sensory-based cognition, but for the present time persons interacting with this animal should be made aware of the importance of the sounds they make, the movements of their bodies, the way they touch the animal, and the odors they emit or carry on their clothing.
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McLean, A. N. (2004). Short-term spatial memory in the domestic horse. Appl. Anim. Behav. Sci., 85(1-2), 93–105.
Abstract: This study investigates the ability of horses to recall a feeding event in a two-point choice apparatus. Twelve horses were individually tested whereby they were maintained immobile in a test arena and visually and aurally experienced the delivery of food into one of two feed goals. The horses were then released to make their choice in two experimental contexts: immediate release after experiencing the delivery of food, and release 10 s after food delivery. Each horse performed 40 immediate-release (IR) trials, followed by forty 10-s release trials over a 3-day period. In addition, the same horses were tested 3 months later in the spring with the same number and sequence of trials. Results were analysed by log-linear analysis of frequencies. Results showed that while horses were able to achieve the correct feed goal choice in the immediate-release trials, they were unsuccessful with the 10-s release trials. This suggests that there are limitations in recall abilities in horses, in that they may not possess a prospective type of memory. There are welfare and training implications in these findings concerning the effects of overestimating the mental abilities of horses during training and the effects of delays in reinforcements.
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Nicol, C. J. (2002). Equine learning: progress and suggestions for future research. Appl. Anim. Behav. Sci., 78(2-4), 193–208.
Abstract: Horses are well able to form classical and instrumental associations and so the focus of much recent research has been on the stimulus control of instrumental learning. Horses appear to discriminate using spatial cues more easily than other stimulus features, as indicated both by the speed of initial task acquisition and by the extent to which acquired discriminations can be reversed. Phenomena associated with discrimination learning in laboratory animals, including generalisation and peak shift, have been demonstrated in horses. However, the ability of horses to classify stimuli into categories is more controversial. Although there is some evidence that horses may be able to form categories based on similarities in the physical appearance of different stimuli, there is currently no evidence that they are able to develop abstract concepts. Their performance on social learning tasks has also been poor. Few correlations are observed between the learning ability of individual horses on different tasks, suggesting that it may not be possible to classify individual horses as `good' or `poor' learners. Better learning performance by horses that are naturally calm is probably due to reduced interference in the learning process. Correct handling procedures can lower reactivity levels in horses, and may facilitate learning in some circumstances. Future research on equine learning needs to take into account the complex nature of equine social interaction. Studies on the effects of stress on learning, and on social and spatial cognition, are also particularly needed.
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Worden, R. P. (1996). Primate social intelligence. Cognit. Sci., 20(4), 579–616.
Abstract: A computational theory of primate social intelligence is proposed in which primates represent social situations internally by discrete symbol structures, called scripts. Three well-defined computational operations on scripts are sufficient to support social learning, planning, and prediction. This gives a formal, predictive model with which to analyse how primate social knowledge is acquired, as well as how it is used. The theory is compared with primate data, such as Cheney and Seyfarth's observations of vervet monkeys. It gives simple, understandable script-based analyses of many observed phenomena--such as the recognition and use of kin relations, learning of alarm calls, habituation to calls, knowledge of rank, tactical deception, and attachment behaviour. I argue that a tight, concise theory of social cognition, such as script theory, is needed to explain the rapid learning and social guile seen in primates. It also has the benefits of simplicity and testability. The extension of scripts to incorporate a primate theory of mind is described in a subsequent paper.
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Plotnik, J. M., de Waal, F. B. M., & Reiss, D. (2006). Self-recognition in an Asian elephant. Proc. Natl. Acad. Sci. U.S.A., 103(45), 17053–17057.
Abstract: Considered an indicator of self-awareness, mirror self-recognition (MSR) has long seemed limited to humans and apes. In both phylogeny and human ontogeny, MSR is thought to correlate with higher forms of empathy and altruistic behavior. Apart from humans and apes, dolphins and elephants are also known for such capacities. After the recent discovery of MSR in dolphins (Tursiops truncatus), elephants thus were the next logical candidate species. We exposed three Asian elephants (Elephas maximus) to a large mirror to investigate their responses. Animals that possess MSR typically progress through four stages of behavior when facing a mirror: (i) social responses, (ii) physical inspection (e.g., looking behind the mirror), (iii) repetitive mirror-testing behavior, and (iv) realization of seeing themselves. Visible marks and invisible sham-marks were applied to the elephants' heads to test whether they would pass the litmus “mark test” for MSR in which an individual spontaneously uses a mirror to touch an otherwise imperceptible mark on its own body. Here, we report a successful MSR elephant study and report striking parallels in the progression of responses to mirrors among apes, dolphins, and elephants. These parallels suggest convergent cognitive evolution most likely related to complex sociality and cooperation.
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Cameron, E. Z. (2004). Facultative adjustment of mammalian sex ratios in support of the Trivers-Willard hypothesis: evidence for a mechanism. Proc Biol Sci, 271(1549), 1723–1728.
Abstract: Evolutionary theory predicts that mothers of different condition should adjust the birth sex ratio of their offspring in relation to future reproductive benefits. Published studies addressing variation in mammalian sex ratios have produced surprisingly contradictory results. Explaining the source of such variation has been a challenge for sex-ratio theory, not least because no mechanism for sex-ratio adjustment is known. I conducted a meta-analysis of previous mammalian sex-ratio studies to determine if there are any overall patterns in sex-ratio variation. The contradictory nature of previous results was confirmed. However, studies that investigated indices of condition around conception show almost unanimous support for the prediction that mothers in good condition bias their litters towards sons. Recent research on the role of glucose in reproductive functioning have shown that excess glucose favours the development of male blastocysts, providing a potential mechanism for sex-ratio variation in relation to maternal condition around conception. Furthermore, many of the conflicting results from studies on sex-ratio adjustment would be explained if glucose levels in utero during early cell division contributed to the determination of offspring sex ratios.
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