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Williams, N. (1997). Evolutionary psychologists look for roots of cognition (Vol. 275).
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Healy, S. D., Braham, S. R., & Braithwaite, V. A. (1999). Spatial working memory in rats: no differences between the sexes. Proc Biol Sci, 266(1435), 2303–2308.
Abstract: In a number of mammalian species, males appear to have superior spatial abilities to females. The favoured explanations for this cognitive difference are hormonal, with higher testosterone levels in males than females leading to better spatial performance, and evolutionary, where sexual selection has favoured males with increased spatial abilities for either better navigational skills in hunting or to enable an increased territory size. However, an alternative explanation for this sex difference focuses on the role of varying levels of oestrogen in females in spatial cognition (the 'fertility and parental care' hypothesis). One possibility is that varying oestrogen levels result in variation in spatial learning and memory so that, when tested across the oestrous cycle, females perform as well as males on days of low oestrogen but more poorly on days of high oestrogen. If day in the oestrous cycle is not taken into account then, across an experiment, any sex differences found would always produce male superiority. We used a spatial working memory task in a Morris water maze to test the spatial learning and memory abilities of male and female rats. The rats were tested across a number of consecutive days during which the females went through four oestrous cycles. We found no overall sex differences in latencies to reach a submerged platform in a Morris water maze but, on the day of oestrus (low oestrogen), females took an extra swim to learn the platform's location (a 100% increase over the other days in the cycle). Female swim speed also varied across the oestrous cycle but females were no less active on the day of oestrus. These results oppose the predictions of the fertility and parental care hypothesis.
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Whishaw, I. Q., Sacrey, L. - A. R., & Gorny, B. (2009). Hind limb stepping over obstacles in the horse guided by place-object memory. Behav. Brain. Res., 198(2), 372–379.
Abstract: An animal that has stepped over an obstacle with its forelimbs uses a memory of the obstacle to guide the hind limbs so that they also clear the obstacle, even in situations in which long pauses are introduced between forelimb and hind limb stepping. To further clarify the features of hind limb obstacle clearance memory, the present study examined hind limb obstacle clearance in the horse. A rider guided horses over obstacles and paused the horse over obstacles in tests that examined the relationship between forelimb and hind limb stepping, with the following results. First, the horses displayed memory for an obstacle as measured by hind limb lifting over the obstacle for durations lasting as long as 15Â min. The response was not dependent upon ongoing visualization of the obstacle, as limb lifting was unaffected by visual occlusion with blinders, a blindfold, or by removing the obstacle during the pause. Second, previous experience of stepping over an obstacle led to pause-related hind limb lifting at the object's previous location even on trials for which there was no obstacle and so no preceding forelimb lifting. Third, whereas a horse would lift its hind limbs to clear two successively presented obstacles, replacing an obstacle before the horse after the forelimbs had cleared the obstacle prevented subsequent hind limb lifting at the obstacle's previous location. Taken together the results show that hind limb obstacle clearance is guided by a place-object memory. The results are discussed in relation to the differential sensory and memonic control of forelimb and hind limb stepping with the suggestion that place-object memory can guide hind stepping as well as overshadow working memory from front leg stepping.
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Rapin, V., Poncet, P. A., Burger, D., Mermod, C., & Richard, M. A. (2007). [Measurement of the attention time in the horse]. Schweiz Arch Tierheilkd, 149(2), 77–83.
Abstract: A study carried out on 49 horses showed that it is possible to measure the attention time by operant conditioning. After teaching horses an instrumental task using a signal, we were then able to test their attention time by asking them to prolong it increasingly while setting success and failure criteria. Two tests were performed 3 weeks apart. The 2nd test was feasible without relearning, a proof of memory, and was repeatable, a proof of consistency in the attention time. A significant difference was observed between the 3 age groups. Young horses often performed very well during the 1st test but their attention dropped in the 2nd test while older horses were more stable with respect to attention and even increased it slightly. The study shows that there are individual differences but it was not possible to prove a significant influence of breed, gender and paternal influence. Consequently, learning appears to be one of the most interesting approaches for evaluating the attention of horses and for observing their behaviour.
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Zentall, T. R. (2002). A cognitive behaviorist approach to the study of animal behavior. J Gen Psychol, 129(4), 328–363.
Abstract: Traditional psychological approaches to animal learning and behavior have involved either the atheoretical behaviorist approach proposed by B. F. Skinner (1938), in which input-output relations are described in response to environmental manipulations, or the theoretical behaviorist approach offered by C. L Hull (1943), in which associations mediated by several hypothetical constructs and intervening variables are formed between stimuli and responses. Recently, the application of a cognitive behaviorist approach to animal learning and behavior has been found to have considerable value as a research tool. This perspective has grown out of E. C. Tolman's cognitive approach to learning in which behavior is mediated by mechanisms that are not directly observable but can be inferred from the results of critical experiments. In the present article, the author presents several examples of the successful application of the cognitive behaviorist approach. In each case, the experiments have been designed to distinguish between more traditional mechanisms and those mediated by hypothesized internal representations. These examples were selected because the evidence suggests that some form of active cognitive organization is needed to account for the behavioral results.
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Gibson, B. M., & Shettleworth, S. J. (2005). Place versus response learning revisited: tests of blocking on the radial maze. Behav Neurosci, 119(2), 567–586.
Abstract: Neurobiological and behavioral research indicates that place learning and response learning occur simultaneously, in parallel. Such findings seem to conflict with theories of associative learning in which different cues compete for learning. The authors conducted place+response training on a radial maze and then tested place learning and response learning separately by reconfiguring the maze in various ways. Consistent with the effects of manipulating place and response systems in the brain (M. G. Packard & J. L. McGaugh, 1996), well-trained rats showed strong place learning and strong response learning. Three experiments using associative blocking paradigms indicated that prior response learning interferes with place learning. Blocking and related tests can be used to better understand how memory systems interact during learning.
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Devenport, J. A., Patterson, M. R., & Devenport, L. D. (2005). Dynamic averaging and foraging decisions in horses (Equus callabus). J. Comp. Psychol., 119(3), 352–358.
Abstract: The variability of most environments taxes foraging decisions by increasing the uncertainty of the information available. One solution to the problem is to use dynamic averaging, as do some granivores and carnivores. Arguably, the same strategy could be useful for grazing herbivores, even though their food renews and is more homogeneously distributed. Horses (Equus callabus) were given choices between variable patches after short or long delays. When patch information was current, horses returned to the patch that was recently best, whereas those without current information matched choices to the long-term average values of the patches. These results demonstrate that a grazing species uses dynamic averaging and indicate that, like granivores and carnivores, they can use temporal weighting to optimize foraging decisions.
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Hausberger, M., Bruderer, C., Le Scolan, N., & Pierre, J. - S. (2004). Interplay between environmental and genetic factors in temperament/personality traits in horses (Equus caballus). J Comp Psychol, 118(4), 434–446.
Abstract: The aim of the present study was to broach the question of the relative influence of different genetic and environmental factors on different temperament/personality traits of horses (Equus caballus). The researchers submitted 702 horses to standardized experimental tests and investigated 9 factors, either genetic or environmental. Genetic factors, such as sire or breed, seemed to influence more neophobic reactions, whereas environmental factors, such as the type of work, seemed to play a more dominant role in reactions to social separation or learning abilities. Additive effects were evident, showing how environmental factors may modulate behavioral traits. This study constitutes a first step toward understanding the relative weights of genetic factors and how the environment may intervene in determining individual behavioral characteristics.
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Roper, K. L., & Zentall, T. R. (1993). Directed forgetting in animals. Psychol Bull, 113(3), 513–532.
Abstract: Directed-forgetting research with animals suggests that animals show disrupted test performance only under certain conditions. Important variables are (a) whether during training, the cue to forget (F cue) signals nonreward (i.e., that the trial is over) versus reward (i.e., that reinforcement can be obtained) and (b) given that reinforcement can be obtained on F-cue trials, whether the post-F-cue response pattern is compatible with the baseline memory task. It is proposed that some findings of directed forgetting can be attributed to trained response biases, whereas others may be attributable perhaps to frustration-produced interference. It is suggested that directed forgetting in animals should be studied using procedures similar to those used to study directed forgetting in humans. This can be accomplished by presenting, within a trial, both to-be-remembered and to-be-forgotten material.
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Healy, S. D., & Jones, C. M. (2002). Animal learning and memory: an integration of cognition and ecology. Zoology, 105(4), 321–327.
Abstract: Summary A wonderfully lucid framework for the ways to understand animal behaviour is that represented by the four [`]whys' proposed by Tinbergen (1963). For much of the past three decades, however, these four avenues have been pursued more or less in parallel. Functional questions, for example, have been addressed by behavioural ecologists, mechanistic questions by psychologists and ethologists, ontogenetic questions by developmental biologists and neuroscientists and phylogenetic questions by evolutionary biologists. More recently, the value of integration between these differing views has become apparent. In this brief review, we concentrate especially on current attempts to integrate mechanistic and functional approaches. Most of our understanding of learning and memory in animals comes from the psychological literature, which tends to use only rats or pigeons, and more occasionally primates, as subjects. The underlying psychological assumption is of general processes that are similar across species and contexts rather than a range of specific abilities. However, this does not seem to be entirely true as several learned behaviours have been described that are specific to particular species or contexts. The first conspicuous exception to the generalist assumption was the demonstration of long delay taste aversion learning in rats (Garcia et al., 1955), in which it was shown that a stimulus need not be temporally contiguous with a response for the animal to make an association between food and illness. Subsequently, a number of other examples, such as imprinting and song learning in birds (e.g., Bolhuis and Honey, 1998; Catchpole and Slater, 1995; Horn, 1998), have been thoroughly researched. Even in these cases, however, it has been typical for only a few species to be studied (domestic chicks provide the [`]model' imprinting species and canaries and zebra finches the song learning [`]models'). As a result, a great deal is understood about the neural underpinnings and development of the behaviour, but substantially less is understood about interspecific variation and whether variation in behaviour is correlated with variation in neural processing (see review by Tramontin and Brenowitz, 2000 but see ten Cate and Vos, 1999).
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