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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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Rizzolatti, G., Fogassi, L., & Gallese, V. (2006). Mirrors of the mind. Sci Am, 295(5), 54–61.
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Brannon, E. M., & Terrace, H. S. (1998). Ordering of the numerosities 1 to 9 by monkeys. Science, 282(5389), 746–749.
Abstract: A fundamental question in cognitive science is whether animals can represent numerosity (a property of a stimulus that is defined by the number of discriminable elements it contains) and use numerical representations computationally. Here, it was shown that rhesus monkeys represent the numerosity of visual stimuli and detect their ordinal disparity. Two monkeys were first trained to respond to exemplars of the numerosities 1 to 4 in an ascending numerical order (1 --> 2 --> 3 --> 4). As a control for non-numerical cues, exemplars were varied with respect to size, shape, and color. The monkeys were later tested, without reward, on their ability to order stimulus pairs composed of the novel numerosities 5 to 9. Both monkeys responded in an ascending order to the novel numerosities. These results show that rhesus monkeys represent the numerosities 1 to 9 on an ordinal scale.
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Zentall, T. R., & Riley, D. A. (2000). Selective attention in animal discrimination learning. J Gen Psychol, 127(1), 45–66.
Abstract: The traditional approach to the study of selective attention in animal discrimination learning has been to ask if animals are capable of the central selective processing of stimuli, such that certain aspects of the discriminative stimuli are partially or wholly ignored while their relationships to each other, or other relevant stimuli, are processed. A notable characteristic of this research has been that procedures involve the acquisition of discriminations, and the issue of concern is whether learning is selectively determined by the stimulus dimension defined by the discriminative stimuli. Although there is support for this kind of selective attention, in many cases, simpler nonattentional accounts are sufficient to explain the results. An alternative approach involves procedures more similar to those used in human information-processing research. When selective attention is studied in humans, it generally involves the steady state performance of tasks for which there is limited time allowed for stimulus input and a relatively large amount of relevant information to be processed; thus, attention must be selective or divided. When this approach is applied to animals and alternative accounts have been ruled out, stronger evidence for selective or divided attention in animals has been found. Similar processes are thought to be involved when animals search more natural environments for targets. Finally, an attempt is made to distinguish these top-down attentional processes from more automatic preattentional processes that have been studied in humans and other animals.
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Hothersall, B., Harris, P., Sörtoft, L., & Nicol, C. J. (2010). Discrimination between conspecific odour samples in the horse (Equus caballus). Appl. Anim. Behav. Sci., 126(1-2), 37–44.
Abstract: Behavioural observations suggest that smell is important in social discriminations between horses but balanced studies of this capacity are lacking. We used a habituation-discrimination procedure to investigate the ability of horses to distinguish between pairs of odour samples from different individuals. In Study 1, separate tests were conducted for urine, faeces or fleece fabric previously rubbed on the coat (to pick up body odour samples (BOS)) and donor pairs differed in sex, and age. 10 pregnant mares each underwent three tests, one per sample type. A test consisted of three successive 2-min presentations of a sample from Individual A with a simultaneous presentation of a sample from Individual B during the final presentation. Doubly repeated measures ANOVA indicated a main effect of sample type on investigative response (df = 2, f = 7.98, P = 0.004): durations were longer for BOS than for urine or faeces but habituation across trials was most consistent for urine. In the final presentation, mares demonstrated discrimination by investigating the novel urine sample (B) more than the repeated sample (novel: median 8.0 s, IQR = 10; repeated: median 2.5 s, IQR = 6; z = -2.558, P = 0.008). In Study 2, urine samples from castrated male donors were used and neither mares nor their 4-month-old foals discriminated between samples from different individuals in the final presentation. The findings suggest that urine odour may contain some information that horses can use to discriminate between conspecifics. This may be limited to the level of broad categories such as sex or reproductive status; further investigation is needed to reveal what functional information can be transmitted and what compounds are involved.
Keywords: Horse; Olfaction; Social discrimination; Urine; Faeces
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Gabor, V., & Gerken, M. (2010). Horses use procedural learning rather than conceptual learning to solve matching to sample. Appl. Anim. Behav. Sci., 126(3-4), 119–124.
Abstract: Research into higher cognitive abilities of the horse may be limited by developing the adequate experimental design. In this study four pony mares between 8 and 19 years old were included. Three of them reached the criterion to be tested in a new design of matching to sample using a black circle and a cross as visual cues attached to an apparatus. The attention was directed to the question of whether the animals are able to concept formation in a given time period or if their decisions depend on other cues or strategies. After familiarization to the testing area and the test procedure, the animals were given 27 sessions of 20 trials each during 14 weeks. While there was no preference for one of the stimuli used, horses showed a significant left sidedness. None of the mares reached the learning criterion of 80% correct answers in one session. However, the ponies showed procedural learning based on correction runs that were given between incorrect decisions, by then selecting the correct stimulus on the other side of the apparatus. This learning type arose in three individuals in session four, six and eleven, respectively. It is concluded that discrimination tasks may be biased by the involvement of unexpected learning strategies, which complicates the interpretation of such tests and may even mask possible conceptualization capabilities.
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Hanggi, E. B., Ingersoll, J. F., & Waggoner, T. L. (2007). Color vision in horses (Equus caballus): deficiencies identified using a pseudoisochromatic plate test. J. Comp. Psychol., 121(1), 65–72.
Abstract: In the past, equine color vision was tested with stimuli composed either of painted cards or photographic slides or through physiological testing using electroretinogram flicker photometry. Some studies produced similar results, but others did not, demonstrating that there was not yet a definitive answer regarding color vision in horses (Equus caballus). In this study, a pseudoisochromatic plate test--which is highly effective in testing color vision both in small children and in adult humans--was used for the first time on a nonhuman animal. Stimuli consisted of different colored dotted circles set against backgrounds of varying dots. The coloration of the circles corresponded to the visual capabilities of different types of color deficiencies (anomalous trichromacy and dichromacy). Four horses were tested on a 2-choice discrimination task. All horses successfully reached criterion for gray circles and demonstration circles. None of the horses were able to discriminate the protan-deutan plate or the individual protan or deutan plates. However, all were able to discriminate the tritan plate. The results suggest that horses are dichromats with color vision capabilities similar to those of humans with red-green color deficiencies.
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Hall, C. A., Cassaday, H. J., Vincent, C. J., & Derrington, A. M. (2006). Cone excitation ratios correlate with color discrimination performance in the horse (Equus caballus). J Comp Psychol, 120(4), 438–448.
Abstract: Six horses (Equus caballus) were trained to discriminate color from grays in a counterbalanced sequence in which lightness cues were irrelevant. Subsequently, the pretrained colors were presented in a different sequence. Two sets of novel colors paired with novel grays were also tested. Performance was just as good in these transfer tests. Once the horse had learned to select the chromatic from the achromatic stimulus, regardless of the specific color, they were immediately able to apply this rule to novel stimuli. In terms of the underlying visual mechanisms, the present study showed for the first time that the spectral sensitivity of horse cone photopigments, measured as cone excitation ratios, was correlated with color discrimination performance, measured as accuracy, repeated errors, and latency of approach.
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Langbein, J., Nurnberg, G., Puppe, B., & Manteuffel, G. (2006). Self-Controlled Visual Discrimination Learning of Group-Housed Dwarf Goats (Capra hircus): Behavioral Strategies and Effects of Relocation on Learning and Memory. J. Comp. Psychol., 120(1), 58–66.
Abstract: In most studies on animal learning, individual animals are tested separately in a specific learning environment and with a limited number of trials per day. An alternative approach is to test animals in a familiar environment in their social group. In this study, the authors--applying a fully automated learning device--investigated voluntary, self-controlled visual shape discrimination learning of group-housed dwarf goats (Capra hircus). The majority of the tested goats showed successful shape discrimination, which indicates the adaptive value of an effective learning strategy. However, in each group, a few individual goats developed behavioral strategies different from shape discrimination to get reward. Relocation impairs memory retrieval (probably by attention shifting) only temporarily for previously learnt shapes. The results demonstrate the usefulness of a self-controlled learning paradigm to assess learning abilities of social species in their normal social settings. This may be especially relevant for captive animals to improve their welfare.
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Sutton, J. E., & Shettleworth, S. J. (2005). Internal sense of direction and landmark use in pigeons (Columba livia). J Comp Psychol, 119(3), 273–284.
Abstract: The relative importance of an internal sense of direction based on inertial cues and landmark piloting for small-scale navigation by White King pigeons (Columba livia) was investigated in an arena search task. Two groups of pigeons differed in whether they had access to visual cues outside the arena. In Experiment 1, pigeons were given experience with 2 different entrances and all pigeons transferred accurate searching to novel entrances. Explicit disorientation before entering did not affect accuracy. In Experiments 2-4, landmarks and inertial cues were put in conflict or tested 1 at a time. Pigeons tended to follow the landmarks in a conflict situation but could use an internal sense of direction to search when landmarks were unavailable.
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