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Griffiths, D. P., & Clayton, N. S. (2001). Testing episodic memory in animals: A new approach. Physiol. Behav., 73(5), 755–762.
Abstract: Episodic memory involves the encoding and storage of memories concerned with unique personal experiences and their subsequent recall, and it has long been the subject of intensive investigation in humans. According to Tulving's classical definition, episodic memory “receives and stores information about temporally dated episodes or events and temporal-spatial relations among these events.” Thus, episodic memory provides information about the `what' and `when' of events (`temporally dated experiences') and about `where' they happened (`temporal-spatial relations'). The storage and subsequent recall of this episodic information was thought to be beyond the memory capabilities of nonhuman animals. Although there are many laboratory procedures for investigating memory for discrete past episodes, until recently there were no previous studies that fully satisfied the criteria of Tulving's definition: they can all be explained in much simpler terms than episodic memory. However, current studies of memory for cache sites in food-storing jays provide an ethologically valid model for testing episodic-like memory in animals, thereby bridging the gap between human and animal studies memory. There is now a pressing need to adapt these experimental tests of episodic memory for other animals. Given the potential power of transgenic and knock-out procedures for investigating the genetic and molecular bases of learning and memory in laboratory rodents, not to mention the wealth of knowledge about the neuroanatomy and neurophysiology of the rodent hippocampus (a brain area heavily implicated in episodic memory), an obvious next step is to develop a rodent model of episodic-like memory based on the food-storing bird paradigm. The development of a rodent model system could make an important contribution to our understanding of the neural, molecular, and behavioral mechanisms of mammalian episodic memory.
Keywords: Episodic memory; Food-caching; Animal models
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Zentall, T. R. (2006). Mental time travel in animals: a challenging question. Behav. Process., 72(2), 173–183.
Abstract: Humans have the ability to mentally recreate past events (using episodic memory) and imagine future events (by planning). The best evidence for such mental time travel is personal and thus subjective. For this reason, it is particularly difficult to study such behavior in animals. There is some indirect evidence, however, that animals have both episodic memory and the ability to plan for the future. When unexpectedly asked to do so, animals can report about their recent past experiences (episodic memory) and they also appear to be able to use the anticipation of a future event as the basis for a present action (planning). Thus, the ability to imagine past and future events may not be uniquely human.
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Zentall, T. R. (1999). Support for a theory of memory for event duration must distinguish between test-trial ambiguity and actual memory loss. J Exp Anal Behav, 72(3), 467–472.
Abstract: Staddon and Higa's (1999) trace-strength theory of timing and memory for event duration can account for pigeons' bias to “choose short” when retention intervals are introduced and to “choose long” when, following training with a fixed retention interval, retention intervals are shortened. However, it does not account for the failure of pigeons to choose short when the intertrial interval is distinct from the retention interval. That finding suggests that stimulus generalization (or ambiguity) between the intertrial interval and the retention interval may result in an effect that has been attributed to memory loss. Such artifacts must be eliminated before a theory of memory for event duration can be adequately tested.
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Zentall, T. R. (2005). Timing, memory for intervals, and memory for untimed stimuli: the role of instructional ambiguity. Behav. Process., 70(3), 209–222.
Abstract: Theories of animal timing have had to account for findings that the memory for the duration of a timed interval appears to be dramatically shorted within a short time of its termination. This finding has led to the subjective shortening hypothesis and it has been proposed to account for the poor memory that animals appear to have for the initial portion of a timed interval when a gap is inserted in the to-be-timed signal. It has also been proposed to account for the poor memory for a relatively long interval that has been discriminated from a shorter interval. I suggest here a simpler account in which ambiguity between the gap or retention interval and the intertrial interval results in resetting the clock, rather than forgetting the interval. The ambiguity hypothesis, together with a signal salience mechanism that determines how quickly the clock is reset at the start of the intertrial interval can account for the results of the reported timing experiments that have used the peak procedure. Furthermore, instructional ambiguity rather than memory loss may account for the results of many animal memory experiments that do not involve memory for time.
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Allcroft, D. J., Tolkamp, B. J., Glasbey, C. A., & Kyriazakis, I. (2004). The importance of `memory' in statistical models for animal feeding behaviour. Behav. Process., 67(1), 99–109.
Abstract: We investigate models for animal feeding behaviour, with the aim of improving understanding of how animals organise their behaviour in the short term. We consider three classes of model: hidden Markov, latent Gaussian and semi-Markov. Each can predict the typical `clustered' feeding behaviour that is generally observed, however they differ in the extent to which `memory' of previous behaviour is allowed to affect future behaviour. The hidden Markov model has `lack of memory', the current behavioural state being dependent on the previous state only. The latent Gaussian model assumes feeding/non-feeding periods to occur by the thresholding of an underlying continuous variable, thereby incorporating some `short-term memory'. The semi-Markov model, by taking into account the duration of time spent in the previous state, can be said to incorporate `longer-term memory'. We fit each of these models to a dataset of cow feeding behaviour. We find the semi-Markov model (longer-term memory) to have the best fit to the data and the hidden Markov model (lack of memory) the worst. We argue that in view of effects of satiety on short-term feeding behaviour of animal species in general, biologically suitable models should allow `memory' to play a role. We conclude that our findings are equally relevant for the analysis of other types of short-term behaviour that are governed by satiety-like principles.
Keywords: Cow; Feeding data; Bouts; Memory; Satiety; Latent structure; Model comparison
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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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Murray, J. K., Singer, E. R., Morgan, K. L., Proudman, C. J., & French, N. P. (2004). Memory decay and performance-related information bias in the reporting of scores by event riders. Preventive Veterinary Medicine, 63(3-4), 173–182.
Abstract: We used data from a case-control study investigating risk factors for horse falls in the cross-country phase of eventing in Great Britain (GB) to examine evidence for memory decay and information bias. Responses to two questions obtained by telephone for 173 cases and 521 controls were examined for evidence of differential reporting according to the respondent's case-control status and performance in the dressage and cross-country phases of competitions. Information bias was found in the accuracy of reporting dressage penalty scores when analysed as a function of performance level (good/poor). Poor dressage performers were less likely to report accurate dressage scores than good performers. The accuracy of reporting dressage scores decreased as the time between the event and questionnaire completion increased, with no case-control interaction. Competitors who incurred cross-country jumping penalties at the event preceding the selected event reported their cross-country scores with less accuracy when compared with competitors who incurred no penalties. No information bias was found when the reporting of dressage and cross-country scores were analysed as a function of respondent category (case/control).
Keywords: Reporting bias; Memory decay; Risk factors; Horse; Cross-country
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Shettleworth, S. J. (2003). Memory and hippocampal specialization in food-storing birds: challenges for research on comparative cognition. Brain Behav Evol, 62(2), 108–116.
Abstract: The three-way association among food-storing behavior, spatial memory, and hippocampal enlargement in some species of birds is widely cited as an example of a new 'cognitive ecology' or 'neuroecology.' Whether this relationship is as strong as it first appears and whether it might be evidence for an adaptive specialization of memory and hippocampus in food-storers have recently been the subject of some controversy [Bolhuis and Macphail, 2001; Macphail and Bolhuis, 2001]. These critiques are based on misconceptions about the nature of adaptive specializations in cognition, misconceptions about the uniformity of results to be expected from applying the comparative method to data from a wide range of species, and a narrow view of what kinds of cognitive adaptations are theoretically interesting. New analyses of why food-storers (black-capped chickadees, Poecile Atricapilla) respond preferentially to spatial over color cues when both are relevant in a memory task show that this reflects a relative superiority of spatial memory as compared to memory for color rather than exceptional spatial attention or spatial discrimination ability. New studies of chickadees from more or less harsh winter climates also support the adaptive specialization hypothesis and suggest that within-species comparisons may be especially valuable for unraveling details of the relationships among ecology, memory, and brain in food-storing species.
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Neuringer, A. (2004). Reinforced variability in animals and people: implications for adaptive action. Am Psychol, 59(9), 891–906.
Abstract: Although reinforcement often leads to repetitive, even stereotyped responding, that is not a necessary outcome. When it depends on variations, reinforcement results in responding that is diverse, novel, indeed unpredictable, with distributions sometimes approaching those of a random process. This article reviews evidence for the powerful and precise control by reinforcement over behavioral variability, evidence obtained from human and animal-model studies, and implications of such control. For example, reinforcement of variability facilitates learning of complex new responses, aids problem solving, and may contribute to creativity. Depression and autism are characterized by abnormally repetitive behaviors, but individuals afflicted with such psychopathologies can learn to vary their behaviors when reinforced for so doing. And reinforced variability may help to solve a basic puzzle concerning the nature of voluntary action.
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Schwartz, B. L., & Evans, S. (2001). Episodic memory in primates. Am. J. Primatol., 55(2), 71–85.
Abstract: Episodic memory refers to a system of memory with the capacity to recollect specific events from an individual's life. Some psychologists have suggested that episodic memory is a uniquely human phenomenon. We challenge that idea and present evidence that great apes and other primates may possess episodic-like memory. We review criteria developed to assess episodic-like memory in nonhumans, and how they apply to primates. In particular, we discuss the criteria of Clayton et al. [2001], who stated that episodic-like memory is based on the retrieval of multiple and integrated components of an event. We then review eight studies examining memory in great apes and apply the Clayton et al. criteria to each of them. We summarize the evidence that is compatible with the existence of episodic-like memory, although none of the data completely satisfy the Clayton et al. criteria. Morover, feelings of pastness and feelings of confidence, which mark episodic memory in humans, have not been empirically addressed in nonhuman primates. Future studies should be directed at these aspects of memory in primates. We speculate on the functional significance of episodic memory in nonhuman primates.
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