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Petherick, J. C., Waddington, D., & Duncan, I. J. H. (1991). Learning to gain access to a foraging and dustbathing substrate by domestic fowl: is `out of sight out of mind'? Behav. Process., 22(3), 213–226.
Abstract: Domestic fowl were deprived of the opportunity to perform litter-related behaviour for three or four days and were tested in a Y-maze (which they had previously been trained to run) for their ability to associate a coloured cue with gaining access to peat. When the goal boxes were within sight of the choice point, most birds chose peat. However, when the birds had to rely solely on the coloured cue only one bird from 12 showed learning. However, the birds seemed to have some expectation of a reward, as they ran faster if, on the previous trial, they had chosen peat. The inability of the birds to learn the association may have been an artefact of the schedule of deprivation and testing, for when they were hungry and tested in the same way they were again unable to learn an association between the same coloured cue and food reward. The experiment with peat was repeated using “massed” trials (several trials in immediate succession) during training and testing and six from 15 birds showed learning. These results suggest that the initial failure to learn was probably due to the training and testing schedule, that access to peat appears to be rewarding and that hens can learn an association between an abstract cue and a rewarding consequence. This is consistent with the possibility that domestic fowls may have some cognitive representation of peat when it is out of sight.
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Pickens, C. L., & Holland, P. C. (2004). Conditioning and cognition. Neurosci Biobehav Rev, 28(7), 651–661.
Abstract: Animals' abilities to use internal representations of absent objects to guide adaptive behavior and acquire new information, and to represent multiple spatial, temporal, and object properties of complex events and event sequences, may underlie many aspects of human perception, memory, and symbolic thought. In this review, two classes of simple associative learning tasks that address these core cognitive capacities are discussed. The first set, including reinforcer revaluation and mediated learning procedures, address the power of Pavlovian conditioned stimuli to gain access, through learning, to representations of upcoming events. The second set of investigations concern the construction of complex stimulus representations, as illustrated in studies of contextual learning, the conjunction of explicit stimulus elements in configural learning procedures, and recent studies of episodic-like memory. The importance of identifying both cognitive process and brain system bases of performance in animal models is emphasized.
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Plotnik, J., Nelson, P. A., & de Waal, F. B. M. (2003). Visual field information in the face perception of chimpanzees (Pan troglodytes). Ann N Y Acad Sci, 1000, 94–98.
Abstract: Evidence for a visual field advantage (VFA) in the face perception of chimpanzees was investigated using a modification of a free-vision task. Four of six chimpanzee subjects previously trained on a computer joystick match-to-sample paradigm were able to distinguish between images of neutral face chimeras consisting of two left sides (LL) or right sides (RR) of the face. While an individual's ability to make this distinction would be unlikely to determine their suitability for the VFA tests, it was important to establish that distinctive information was available in test images. Data were then recorded on their choice of the LL vs. RR chimera as a match to the true, neutral image; a bias for one of these options would indicate an hemispatial visual field advantage. Results suggest that chimpanzees, unlike humans, do not exhibit a left visual field advantage. These results have important implications for studies on laterality and asymmetry in facial signals and their perception in primates.
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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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Poti, P. (2005). Chimpanzees' constructional praxis (Pan paniscus, P. troglodytes). Primates, 46(2), 103–113.
Abstract: This study investigated chimpanzees' spontaneous spatial constructions with objects and especially their ability to repeat inter-object spatial relations, which is basic to understanding spatial relations at a higher level than perception or recognition. Subjects were six chimpanzees-four chimpanzees and two bonobos-aged 6-21 years, all raised in a human environment from an early age. Only minor species differences, but considerable individual differences were found. The effect of different object samples was assessed through a comparison with a previous study. A common overall chimpanzee pattern was also found. Chimpanzees repeated different types of inter-object spatial relations such as insertion (I), or vertical (V), or next-to (H) relations. However chimpanzees repeated I or V relations with more advanced procedures than when repeating H relations. Moreover, chimpanzees never repeated combined HV relations. Compared with children, chimpanzees showed a specific difficulty in repeating H relations. Repeating H relations is crucial for representing and understanding multiple reciprocal spatial relations between detached elements and for coordinating independent positions in space. Therefore, the chimpanzees' difficulty indicates a fundamental difference in constructive space in comparison to humans. The findings are discussed in relation to issues of spatial cognition and tool use.
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Proops, L., McComb, K., & Reby, D. (2008). Cross-modal individual vocal recognition in the domestic horse. In IESM 2008.
Abstract: Horses fulfill many of the criteria for a species in which it would be adaptive to be capable of individual recognition: they are highly social, form strong and long lasting bonds, their affiliations are rarely kin based, they have a fission-fusion social structure and they possess inter and intra-group dominance hierarchies.
We used a novel cross-modal, expectancy violation paradigm to provide the first systematic evidence that a non-human animal – the domestic horse- is capable of cross modal recognition. We believe this paradigm could provide an ideal way to study individual recognition across a wide range of species.
For full published details see: Proops L, McComb K, Reby D (2009) Cross-modal individual recognition in domestic horses (Equus caballus). Proc Natl Acad Sci U S A 106: 947-951.
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Proops, L., McComb, K., & Reby, D. (2008). Horse-human interactions: Attention attribution and the use of human cues by domestic horses (Equus caballus). In IESM 2008.
Abstract: Recent research has shown that domestic dogs are particularly good at reading human attentional cues, often outperforming chimpanzees and hand reared wolves [1, 2]. It has been suggested that the close evolutionary relationship between humans and dogs has led to the development of this ability, however very few other species have been studied [3]. We tested the ability of 24 domestic horses to discriminate between an attentive and inattentive person when choosing whom to approach for food. While the attentive person faced forwards, the inattentive person either stood with their body turned 180° away from the subject (body orientation condition), stood with their body facing forwards but their head facing away (head orientation condition) or stood facing forwards but with their eyes closed (eyes closed condition). A fourth, mixed condition was included where the attentive person stood with their body facing away from the subjects but their head turned towards the subject while the inattentive person stood with their body facing the subject but their head turned away. Horses chose the attentive person significantly more often using the body cue (n = 24, k = 19, p = 0.003), the head cue (n = 24, k = 18, p = 0.011), and the eye cue (n = 24, k = 19, p = 0.003) but not the mixed cue (n = 24, k = 13, p = 0.42). In an additional pilot study, horses were tested in an object choice task. A human experimenter cued one of two buckets by either tapping the bucket (tap condition), orienting their body towards the bucket and pointing (body and point condition), pointing while facing forwards (point condition) or orienting their body towards the bucket (body condition). If the subjects chose the correct bucket they were rewarded. Subjects were able to use the tap cue (n = 31, k = 21, p = 0.035), body + point cue (n= 31, k = 21, p = 0.035) and the point cue (n = 30, k = 21, p = 0.021) but not the body cue (n = 31, k = 11, p = 0.076). These results taken together suggest that domestic horses are also very sensitive to human attentional cues, including gaze.
Keywords:
social cognition, animal-human interaction, horses, attention attribution, domestication
1. Hare, B., Brown, M., Williamson, C., and Tomasello, M. (2002). The domestication of social cognition in dogs. Science 298, 1634-1636.
2. Gacsi, M., Miklosi, A., Varga, O., Topal, J., and Csanyi, V. (2004). Are readers of our face readers of our minds` Dogs (Canis familiaris) show situation-dependent recognition of human’s attention. Animal Cognition 7, 144-153.
3. Hare, B., and Tomasello, M. (2005). Human-like social skills in dogs? Trends Cogn. Sci. 9, 439-444.
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Proops, L., McComb, K., & Reby, D. (2009). Cross-modal individual recognition in domestic horses (Equus caballus). Proc. Natl. Acad. Sci. U.S.A., 106(3), 947–951.
Abstract: Individual recognition is considered a complex process and, although it is believed to be widespread across animal taxa, the cognitive mechanisms underlying this ability are poorly understood. An essential feature of individual recognition in humans is that it is cross-modal, allowing the matching of current sensory cues to identity with stored information about that specific individual from other modalities. Here, we use a cross-modal expectancy violation paradigm to provide a clear and systematic demonstration of cross-modal individual recognition in a nonhuman animal: the domestic horse. Subjects watched a herd member being led past them before the individual went of view, and a call from that or a different associate was played from a loudspeaker positioned close to the point of disappearance. When horses were shown one associate and then the call of a different associate was played, they responded more quickly and looked significantly longer in the direction of the call than when the call matched the herd member just seen, an indication that the incongruent combination violated their expectations. Thus, horses appear to possess a cross-modal representation of known individuals containing unique auditory and visual/olfactory information. Our paradigm could provide a powerful way to study individual recognition across a wide range of species.
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Proops, L., Walton, M., & McComb, K. (2010). The use of human-given cues by domestic horses, Equus caballus, during an object choice task. Anim. Behav., 79(6), 1205–1209.
Abstract: Selection pressures during domestication are thought to lead to an enhanced ability to use human-given cues. Horses fulfil a wide variety of roles for humans and have been domesticated for at least 5000 years but their ability to read human cues has not been widely studied. We tested the ability of 28 horses to attend to human-given cues in an object choice task. We included five different cues: distal sustained pointing, momentary tapping, marker placement, body orientation and gaze (head) alternation. Horses were able to use the pointing and marker placement cues spontaneously but not the tapping, body orientation and gaze alternation cues. The overall pattern of responding suggests that horses may use cues that provide stimulus enhancement at the time of choice and do not have an understanding of the communicative nature of the cues given. As such, their proficiency at this task appears to be inferior to that of domestic dogs, Canis lupus familiaris, but similar to that of domestic goats, Caprus hircus.
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Real, L. A. (1991). Animal choice behavior and the evolution of cognitive architecture. Science, 253(5023), 980–986.
Abstract: Animals process sensory information according to specific computational rules and, subsequently, form representations of their environments that form the basis for decisions and choices. The specific computational rules used by organisms will often be evolutionarily adaptive by generating higher probabilities of survival, reproduction, and resource acquisition. Experiments with enclosed colonies of bumblebees constrained to foraging on artificial flowers suggest that the bumblebee's cognitive architecture is designed to efficiently exploit floral resources from spatially structured environments given limits on memory and the neuronal processing of information. A non-linear relationship between the biomechanics of nectar extraction and rates of net energetic gain by individual bees may account for sensitivities to both the arithmetic mean and variance in reward distributions in flowers. Heuristic rules that lead to efficient resource exploitation may also lead to subjective misperception of likelihoods. Subjective probability formation may then be viewed as a problem in pattern recognition subject to specific sampling schemes and memory constraints.
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