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Scordato, E. S., & Drea, C. M. (2007). Scents and sensibility: information content of olfactory signals in the ringtailed lemur, Lemur catta. Anim. Behav., 73(2), 301–314.
Abstract: The function of olfactory signalling in social species is less well understood than in asocial species. Consequently, we examined olfactory communication in the ringtailed lemur, a socially complex primate that retains a functional vomeronasal organ, has well-developed scent glands and shows a suite of scent-marking behaviour. To assess the information content of different types of scent gland secretions, we decoupled olfactory cues from the visual and behavioural modalities with which scent marking is normally associated. We presented male and female subjects (signal receivers) with a series of choice tests between odours derived from conspecific donors (signal senders) varying by sex, age, social status and reproductive condition. We additionally examined the influence of the receivers' reproductive state and familiarity with the signaller. The reproductive condition, social status and familiarity of senders and receivers affected signal transmission; specifically, male receivers attended most to the odours of conspecifics in breeding condition and to the odours of familiar, dominant animals. By contrast, females varied their responses according to both their own reproductive state and that of the sender. Based on male and female patterns of countermarking, we suggest that scent marking serves a function in intergroup spacing and intrasexual competition for both sexes, as might be expected in a female-dominant species. By contrast, minimal female interest in male odours counters a female mate choice function for scent marking in this species. Nevertheless, scent marks are critical to male-male competition and, therefore, may be subject to sexual selection.
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Dreier, S., van Zweden, J. S., & D'Ettorre, P. (2007). Long-term memory of individual identity in ant queens. Biol Lett, 3(5), 459–462.
Abstract: Remembering individual identities is part of our own everyday social life. Surprisingly, this ability has recently been shown in two social insects. While paper wasps recognize each other individually through their facial markings, the ant, Pachycondyla villosa, uses chemical cues. In both species, individual recognition is adaptive since it facilitates the maintenance of stable dominance hierarchies among individuals, and thus reduces the cost of conflict within these small societies. Here, we investigated individual recognition in Pachycondyla ants by quantifying the level of aggression between pairs of familiar or unfamiliar queens over time. We show that unrelated founding queens of P. villosa and Pachycondyla inversa store information on the individual identity of other queens and can retrieve it from memory after 24h of separation. Thus, we have documented for the first time that long-term memory of individual identity is present and functional in ants. This novel finding represents an advance in our understanding of the mechanism determining the evolution of cooperation among unrelated individuals.
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Tibbetts, E. A. (2002). Visual signals of individual identity in the wasp Polistes fuscatus. Proc. Roy. Soc. Lond. B Biol. Sci., 269(1423), 1423–1428.
Abstract: Individual recognition is an essential component of interactions in many social systems, but insects are often thought incapable of the sophistication necessary to recognize individuals. If this were true, it would impose limits on the societies that insects could form. For example, queens and workers of the paper wasp Polistes fuscatus form a linear dominance hierarchy that determines how food, work and reproduction are divided within the colony. Such a stable hierarchy would be facilitated if individuals of different ranks have some degree of recognition. P. fuscatus wasps have, to our knowledge, previously undocumented variability in their yellow facial and abdominal markings that are intriguing candidates for signals of individual identity. Here, I describe these highly variable markings and experimentally test whether P. fuscatus queens and workers use these markings to identify individual nest-mates visually. I demonstrate that individuals whose yellow markings are experimentally altered with paint receive more aggression than control wasps who are painted in a way that does not alter their markings. Further, aggression declines towards wasps with experimentally altered markings as these novel markings become familiar to their nestmates. This evidence for individual recognition in P. fuscatus indicates that interactions between insects may be even more complex than previously anticipated.
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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.
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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.
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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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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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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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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.
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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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