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Lim, M. M., & Young, L. J. (2006). Neuropeptidergic regulation of affiliative behavior and social bonding in animals. Hormon. Behav., 50(4), 506–517.
Abstract: Social relationships are essential for maintaining human mental health, yet little is known about the brain mechanisms involved in the development and maintenance of social bonds. Animal models are powerful tools for investigating the neurobiological mechanisms regulating the cognitive processes leading to the development of social relationships and for potentially extending our understanding of the human condition. In this review, we discuss the roles of the neuropeptides oxytocin and vasopressin in the regulation of social bonding as well as related social behaviors which culminate in the formation of social relationships in animal models. The formation of social bonds is a hierarchical process involving social motivation and approach, the processing of social stimuli and formation of social memories, and the social attachment itself. Oxytocin and vasopressin have been implicated in each of these processes. Specifically, these peptides facilitate social affiliation and parental nurturing behavior, are essential for social recognition in rodents, and are involved in the formation of selective mother-infant bonds in sheep and pair bonds in monogamous voles. The convergence of evidence from these animal studies makes oxytocin and vasopressin attractive candidates for the neural modulation of human social relationships as well as potential therapeutic targets for the treatment of psychiatric disorders associated with disruptions in social behavior, including autism.
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Lampe, J. F., & Andre, J. (2012). Cross-modal recognition of human individuals in domestic horses (Equus caballus). Animal Cognition, 15(4), 623–630.
Abstract: This study has shown that domestic horses are capable of cross-modal recognition of familiar humans. It was demonstrated that horses are able to discriminate between the voices of a familiar and an unfamiliar human without seeing or smelling them at the same moment. Conversely, they were able to discriminate the same persons when only exposed to their visual and olfactory cues, without being stimulated by their voices. A cross-modal expectancy violation setup was employed; subjects were exposed both to trials with incongruent auditory and visual/olfactory identity cues and trials with congruent cues. It was found that subjects responded more quickly, longer and more often in incongruent trials, exhibiting heightened interest in unmatched cues of identity. This suggests that the equine brain is able to integrate multisensory identity cues from a familiar human into a person representation that allows the brain, when deprived of one or two senses, to maintain recognition of this person.
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Owen, H., Hall, C., Hallam, S., & Smith, E. (2012). The use of GPS to measure feeding behaviour and activity patterns in the horse (Equus caballus). In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: The global positioning system (GPS) has been used to record activity and monitor habitat use in many animal species. In the horse (Equus caballus) the monitoring of activity and feeding patterns has been used to assess the impact of environmental factors on behaviour and welfare. In free-ranging animals GPS can provide such information but the accuracy and reliability of these devices has yet to be confirmed. The aim of this study was: 1) to compare the results of visual observation with GPS recordings of the horse’s head and neck position (head up (HU) and down (HD)) used to quantify time spent grazing; 2) to test the use of GPS collars to monitor activity patterns where distance, speed and location paths were recorded. In both studies two animals were fitted with Lotek GPS 3300S collars (with integrated GPS data logger and removable battery pack) round the top of the neck. In study 1 two horses were fitted with collars and turned loose into a 20x40m sand arena for 45 minutes. Feed balls and hay were provided (in nets and on the ground) to encourage movement and feeding behaviour for comparison using the two methods (observation from digital video recordings and GPS). HD was recorded by the GPS collars for a significantly longer time (interpreted as feeding/grazing time) than that recorded by observation (p=0.004). However when the visual observation was split into HU, HD and also head in mid-way position (HMW), where the nose of the horse was level or just above the chest, then no difference between the collar (HU and HD) and visual observation for (HU and HD+HMW) was found. It is likely that when in HMW the GPS collar may not be sufficiently angled to trigger the sensor to record HU or the collar may move on the neck. Conclusions relating to time spent feeding should be treated with caution. In study 2, the collars were fitted to two ponies with access to 2.02 hectares of lowland grazing. Activity (distance travelled and speed) and location was recorded for 2 days. The total distance travelled by the ponies in 24 hours (2.84km) and their average speed (4.04m/minute) was calculated and showed no significant difference between day and night. The total area was split into four equal segments and there was no significant difference in the time the ponies spent in each area although they were found to move at slower speeds and stand for longer in some areas. Movement paths could be identified by inputting the GPS collar data into ArcGIS and viewed on Google Maps. There was a high level of comparability observed between the two ponies confirming behavioural synchronicity. As in other species, the use of GPS collars to monitor the movement and location of horses/ponies was found to be effective, but data relating to head position did not provide a reliable means of recording the time spent feeding.
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Hall, C., Rigg, V., Truswell, M., & Owen, H. (2012). Picture recognition of con-specifics and facial expression in the horse (Equus caballus). In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: The management of the domestic horse often requires them to be kept in isolation from con-specifics. Installing a picture of a horse (generally head and neck view) with a view to providing surrogate companionship has been shown to reduce the negative impact of this isolation. This study aimed firstly to compare the spontaneous response of horses (N=10) to a 2-D image of a horse’s face (FP) with their response to a comparable abstract 2-D image (AP). Secondly, the spontaneous response of horses (N=20) to a 2-D image of a horse’s face with the ears forward (PFP positive) was compared with the response to a 2-D image of a horse’s face with the ears back (NFP negative). The posters were A1 sized and displayed in the horse’s own stable. In study 1, one poster was displayed for 5 minutes and the horse’s behaviour video-recorded. This was removed and the second poster was displayed for 5 minutes and the behaviour video-recorded. FP was displayed first for 5 of the horses and AP displayed first for the other 5. The video footage was observed and the behaviour of the horses and number of times they touched the poster recorded. For the purpose of identifying the area of the poster that was touched by the horse it was divided into 4 equal quarters (TL, TR, BL, BR). In FP the nose of the horse in the 2-D image was located in BL, eyes and ears in TL, chest and lower neck in BR and upper neck in TR. In AP each area contained similar but unique abstract patterns of comparable colour to FP. Differences in behaviour were found according to which poster was displayed. FP was touched significantly more than AP (p=0.001) and was looked at more often (p=0.008). With FP the horses spent significantly longer with their ears forward (p=0.008) and licking and chewing (p=0.016). When the number of touches per poster area was compared (FP and AP) a significant difference was found in the number of times that BL (nose) and BR (chest/lower neck) were touched (p=0.011). Both areas were touched more frequently on FP, with BL being touched the most. In study 2 the same experimental protocol was used to compare responses to positive (PFP) or negative (NFP) 2-D images of a horse’s face (same horse in both PFP and NFP). Again, differences in behaviour were found in response to the two posters. PFP was touched significantly more than NFP (p=0.002) and on both posters the area BL (nose) was touched more frequently than the other areas (PFP: p=0.02, NFP: p=0.01). More ears back behaviour (p<0.001) and more ear locked on behaviour (p=0.008) was shown with NFP. The results of these studies indicate that horses can recognize 2-D images as con-specifics as well as responding to differences in facial expression. There is now the potential for further investigation into the importance of other visual cues in recognition and social interaction as well as the application of findings to enhance equine welfare.
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Christensen, J. W., Zharkikh, T., & Chovaux, E. (2011). Object recognition and generalisation during habituation in horses. Appl. Anim. Behav. Sci., 129(2-4), 83–91.
Abstract: The ability of horses to habituate to frightening stimuli greatly increases safety in the horse-human relationship. A recent experiment suggested, however, that habituation to frightening visual stimuli is relatively stimulus-specific in horses and that shape and colour are important factors for object generalisation (Christensen et al., 2008). In a series of experiments, we aimed to further explore the ability of horses (n = 30, 1 and 2-year-old mares) to recognise and generalise between objects during habituation. TEST horses (n = 15) were habituated to a complex object, composed of five simple objects of varying shape and colour, whereas CONTROL horses (n = 15) were habituated to the test arena, but not to the complex object. In the first experiment, we investigated whether TEST horses subsequently reacted less to i) simple objects that were previously part of the complex object (i.e. testing for object recognition) and ii) a novel object (new shape and colour, i.e. testing for object generalisation), compared to CONTROLS. In the second experiment we investigated whether TEST horses reacted to a change in object order and object location. Behavioural reactions to the object, latency to eat, total eating time and heart rate were recorded. Compared to CONTROLS, TEST horses reacted significantly less towards objects, which were previously part of the complex object (e.g. mean heart rate; P = 0.006), indicating object recognition. In contrast to our expectations, TEST horses also reacted significantly less towards the novel object (e.g. mean heart rate; P = 0.018), suggesting that they were capable of object generalisation. We also found that TEST horses showed an increase in exploratory behaviour when objects within the complex object changed order and location (both P < 0.001), whereas there was no increase in heart rate, indicating that the horses were not frightened by the changes. The results demonstrate that it is possible to increase object generalisation in horses by habituating them to a range of colours and shapes simultaneously. This knowledge greatly affects the way in which horses may be trained to react calmly towards frightening objects.
Keywords: Horse; Habituation; Object recognition; Generalisation
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Thor, D. H., & Holloway, W. R. (1982). Social memory of the male laboratory rat. J. Comp. Physiol. Psychol., 96(6), 1000–1006.
Abstract: Used duration of social-investigatory behavior by 36 mature male Long-Evans rats as a measure of individual recognition in 5 experiments to assess social memory. In Exp I, the duration of social investigation during a 2nd exposure to the same juvenile (n[en space]=[en space]12) was directly related to the length of the interexposure interval. In Exp II, Ss were exposed to the same or different juvenile 10 min after an initial 5-min exposure to a novel juvenile; reexposure to the same juvenile elicited significantly less social investigation than an exposure to a different juvenile. Exps III and IV demonstrated that following a 5-min introductory exposure, social memory of the juvenile was relatively brief in comparison with that of mature Ss. Exp V revealed a retroactive interference effect on recently acquired memory for an individual: 12 mature Ss exposed to interpolated social experience engaged in significantly longer investigation of a juvenile than those with no interpolated social experience. The combined results suggest that (1) the rat normally engages in spontaneous learning of individual identity and (2) social memory may be a significant aspect of complex social interactions. (16 ref) (PsycINFO Database Record (c) 2006 APA, all rights reserved)
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Soproni, K., Miklósi, A., Topál, J., & Csányi, V. (2001). Comprehension of human communicative signs in pet dogs (Canis familiaris). J Comp Psychol, 115(2), 122–126.
Abstract: On the basis of a study by D. J. Povinelli, D. T. Bierschwale, and C. G. Cech (1999), the performance of family dogs (Canis familiaris) was examined in a 2-way food choice task in which 4 types of directional cues were given by the experimenter: pointing and gazing, head-nodding (“at target”), head turning above the correct container (“above target”), and glancing only (“eyes only”). The results showed that the performance of the dogs resembled more closely that of the children in D. J. Povinelli et al.'s study, in contrast to the chimpanzees' performance in the same study. It seems that dogs, like children, interpret the test situation as being a form of communication. The hypothesis is that this similarity is attributable to the social experience and acquired social routines in dogs because they spend more time in close contact with humans than apes do, and as a result dogs are probably more experienced in the recognition of human gestures.
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Soproni, K., Miklósi, Á., Topál, J., & Csányi, V. (2002). Dogs' (Canis familiaris) responsiveness to human pointing gestures. J Comp Psychol, 116(1), 27–34.
Abstract: In a series of 3 experiments, dogs (Canis familiaris) were presented with variations of the human pointing gesture: gestures with reversed direction of movement, cross-pointing, and different arm extensions. Dogs performed at above chance level if they could see the hand (and index finger) protruding from the human body contour. If these minimum requirements were not accessible, dogs still could rely on the body position of the signaler. The direction of movement of the pointing arm did not influence the performance. In summary, these observations suggest that dogs are able to rely on relatively novel gestural forms of the human communicative pointing gesture and that they are able to comprehend to some extent the referential nature of human pointing.
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Virányi, Z., Topál, J., Gácsi, M., Miklósi, Á., & Csányi, V. (2004). Dogs respond appropriately to cues of humans' attentional focus. Behav. Process., 66(2), 161–172.
Abstract: Dogs' ability to recognise cues of human visual attention was studied in different experiments. Study 1 was designed to test the dogs' responsiveness to their owner's tape-recorded verbal commands (Down!) while the Instructor (who was the owner of the dog) was facing either the dog or a human partner or none of them, or was visually separated from the dog. Results show that dogs were more ready to follow the command if the Instructor attended them during instruction compared to situations when the Instructor faced the human partner or was out of sight of the dog. Importantly, however, dogs showed intermediate performance when the Instructor was orienting into 'empty space' during the re-played verbal commands. This suggests that dogs are able to differentiate the focus of human attention. In Study 2 the same dogs were offered the possibility to beg for food from two unfamiliar humans whose visual attention (i.e. facing the dog or turning away) was systematically varied. The dogs' preference for choosing the attentive person shows that dogs are capable of using visual cues of attention to evaluate the human actors' responsiveness to solicit food-sharing. The dogs' ability to understand the communicatory nature of the situations is discussed in terms of their social cognitive skills and unique evolutionary history.
Keywords: Animals; *Attention; Bonding, Human-Pet; Communication; *Cues; Dogs; Humans; Recognition (Psychology)
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De Boyer Des Roches, A., Richard-Yris, M. - A., Henry, S., Ezzaouia, M., & Hausberger, M. (2008). Laterality and emotions: visual laterality in the domestic horse (Equus caballus) differs with objects' emotional value. Physiol. Behav., 94(3), 487–490.
Abstract: Lateralization of emotions has received great attention in the last decades, both in humans and animals, but little interest has been given to side bias in perceptual processing. Here, we investigated the influence of the emotional valence of stimuli on visual and olfactory explorations by horses, a large mammalian species with two large monocular visual fields and almost complete decussation of optic fibres. We confronted 38 Arab mares to three objects with either a positive, negative or neutral emotional valence (novel object). The results revealed a gradient of exploration of the 3 objects according to their emotional value and a clear asymmetry in visual exploration. When exploring the novel object, mares used preferentially their right eyes, while they showed a slight tendency to use their left eyes for the negative object. No asymmetry was evidenced for the object with the positive valence. A trend for an asymmetry in olfactory investigation was also observed. Our data confirm the role of the left hemisphere in assessing novelty in horses like in many vertebrate species and the possible role of the right hemisphere in processing negative emotional responses. Our findings also suggest the importance of both hemispheres in the processing positive emotions. This study is, to our knowledge, the first to demonstrate clearly that the emotional valence of a stimulus induces a specific visual lateralization pattern.
Keywords: Animals; Animals, Newborn; Behavior, Animal/physiology; Dominance, Ocular/*physiology; *Emotions; Exploratory Behavior/physiology; Female; Horses/*physiology; Olfactory Pathways/physiology; Pattern Recognition, Visual/*physiology; Photic Stimulation; Pregnancy; Statistics, Nonparametric; Visual Fields/physiology
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