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Cameron, E. Z., Linklater, W. L., Stafford, K. J., & Minot, E. O. (2008). Maternal investment results in better foal condition through increased play behaviour in horses. Anim. Behav., 76(5), 1511–1518.
Abstract: Play behaviour is widespread in mammals, but benefits to play have been difficult to demonstrate. Physical training is one of the many proposed hypotheses, suggesting that males and females should play differently, that increased maternal investment should lead to increases in play, and that increases in play should result in physical advantages. In a population of feral horses, Equus caballus, males and females did not differ in their play behaviour except that males initiated more of their play bouts than females. Maternal condition influenced play behaviour only in males, with sons of mothers in good condition playing more. However, when we controlled for maternal effects by comparing a son and a daughter of the same mother, daughters played more when their mother was in poor condition and sons played more when their mother was in good condition. Mothers of foals that played more lost more condition. Therefore, the difference in play behaviour could not be explained by offspring sex or maternal condition alone, but play behaviour mirrored variation in maternal investment. In addition, those individuals that played more survived better and had better body condition as yearlings despite weaning earlier. Since increased activity has been linked to enhanced musculoskeletal development in domestic horses, we suggest that play provides a link between increased maternal investment, increased body condition and future reproductive success in feral horses, and probably in other species.
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Ward, C., Trisko, R., & Smuts, B. B. (2009). Third-party interventions in dyadic play between littermates of domestic dogs, Canis lupus familiaris. Anim. Behav., 78(5), 1153–1160.
Abstract: Interventions occur when animals interfere in competitive interactions between two or more individuals. Interveners can alter the nature of the ongoing interaction by targeting one party (attacking, biting) and supporting the other. Three theories have been proposed to account for intervention behaviour: kin selection, reciprocity and direct benefits. The kin selection hypothesis predicts that interveners will selectively support relatives over nonrelatives; the reciprocity hypothesis predicts that when intervener [`]A' supports individual [`]B', later [`]B' will intervene and support [`]A'; and the direct benefits hypothesis predicts that target/support patterns should serve the immediate interests of the intervener. We tested the reciprocity and direct benefits hypotheses by exploring third-party interventions in play fighting among littermates of domestic dogs. Interveners in dyadic play did not preferentially target or support preferred playmates of the intervener. Interveners targeted the dog in the losing role at the time of the intervention, and they did not show reciprocity in support. Taken together, these last two findings suggest that littermates benefit directly and use interventions opportunistically to practise offence behaviours directed at littermates already behaving subordinately. Opportunities to practise targeting in a playful setting may help structure dominance relationships among littermates. Additionally, the tendency for puppies to do what the other is doing (target the dog in the losing role) may pave the way for synchronizing cooperative behaviours during group hunting and territorial defence. The types of behaviours used to intervene changed over development, but the outcome following an intervention remained stable.
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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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de Waal, F. B., & Johanowicz, D. L. (1993). Modification of reconciliation behavior through social experience: an experiment with two macaque species. Child Dev, 64(3), 897–908.
Abstract: Reconciliation, defined as a friendly reunion between former opponents shortly after an aggressive encounter, is common in the stumptail macaque (Macaca arctoides) but rare in the rhesus macaque (M. mulatta). Juveniles of the two species were cohoused for 5 months, after which they were observed with conspecifics only. Control rhesus monkeys, matched in age and sex to the experimental subjects, went through the same procedure without exposure to the other species. A threefold increase in the proportion of reconciled fights was measured in the rhesus subjects. The difference emerged gradually during cohousing with the tutor species and was sustained following removal of this species. Other behavior, such as grooming and aggression, decreased over time. It is suggested that the social attitude of the subjects was affected through contact with a species characterized by a more relaxed dominance style.
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Palagi, E. (2008). Sharing the motivation to play: the use of signals in adult bonobos. Anim. Behav., 75(3), 887–896.
Abstract: Gestures and facial displays are involved in regulating many aspects of mammal social life such as aggression, dominance-subordinate relationships, appeasement and play. Playful activity is an interesting behaviour for examining the role of signals as intentional communication systems. When animals play they perform patterns that are used in other serious contexts. To avoid miscommunication, many species have evolved signals to maintain a playful mood. Bonobos, Pan paniscus, with their flexible social relationships and playful propensity, may represent a good model species to test some hypotheses on adult play signalling. I analysed the potential roles of facial play expressions and solitary play in soliciting and regulating social play and found that adult bonobos used the play face (relaxed open-mouth display) in a selective manner. Play faces were more frequent during social than solitary play and, within social play, polyadic sessions (even though less frequent than dyadic sessions) were characterized by a higher frequency of signals. Following the rule of play intensity matching, play faces were more frequent when the two players matched in age and size (sessions among adults). Moreover, among dyads there was a positive correlation between the frequency of aggressive interactions performed and the frequency of play signals used, thus suggesting that signals are crucial in play negotiations among individuals showing high baseline levels of aggression. Finally, solitary play, especially when it involved pirouettes and somersaults, had an important role in triggering social play.
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Beaver, B. V. (1986). Aggressive behavior problems. Vet Clin North Am Equine Pract, 2(3), 635–644.
Abstract: Accurate diagnosis of the cause of aggression in horses is essential to determining the appropriate course of action. The affective forms of aggression include fear-induced, pain-induced, intermale, dominance, protective, maternal, learned, and redirected aggressions. Non-affective aggression includes play and sex-related forms. Irritable aggression and hypertestosteronism in mares are medical problems, whereas genetic factors, brain dysfunction, and self-mutilation are also concerns.
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Nelson, X. J., & Fijn, N. (2013). The use of visual media as a tool for investigating animal behaviour. Animal Behaviour, 85(3), 525–536.
Abstract: In this essay we outline how video-related technology can be used as a tool for studying animal behaviour. We review particular aspects of novel, innovative animal behaviour uploaded by the general public via video-based media on the internet (using YouTube as a specific example). The behaviour of animals, particularly the play behaviour focused on here, is viewed by huge audiences. In this essay we focused on three different kinds of media clips: (1) interspecies play between dogs and a range of other species; (2) object play in horses; and (3) animal responses to stimuli presented on iPads, iPods and iPhones. We argue that the use of video is a good means of capturing uncommon or previously unknown behaviour, providing evidence that these behaviours occur. Furthermore, some of the behaviours featured on YouTube provide valuable insights for future directions in animal behaviour research. If we also take this opportunity to convey our knowledge to a public that seems to be fundamentally interested in animal behaviour, this is a good means of bridging the gap between knowledge among an academic few and the general public.
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Hanggi, E. B., & Ingersoll, J. F. (2009). Long-term memory for categories and concepts in horses (Equus caballus). Anim. Cogn., 13(3), 451–462.
Abstract: Three horses (Equus caballus) with a history of performing cognitive tasks including discrimination learning, categorization, and concept use were tested to evaluate their long-term memory (LTM) in three experiments. In addition, use of LCD multi-displays for stimulus presentation was incorporated into cognition testing protocol for the first time with horses. Experiment 1 tested LTM for discrimination learning that originally occurred 6 years earlier. Five sets of stimuli were used and the two horses tested showed no decrement in performance on four of the sets; however, both horses did score below chance on one set. Experiment 2 examined long-term categorization recall 10 years after horses had demonstrated the ability to make stimulus selections based on shared characteristics within a given category. The horse tested for LTM after the decade-long interval immediately and consistently applied the previously learned categorization rule to not only familiar but also novel sets of stimuli. Experiment 3 tested another horse for LTM for a relative size concept. This horse had originally demonstrated concept rule use in order to select stimuli based on their relative size to one another. More than 7 years later and without further training, this horse reliably applied the previously established size concept to both familiar and novel sets of stimuli. These findings are the first reports of long-term categorical and conceptual memory in horses and are consistent with observations of domestic and wild horses, which indicate that behavioral and ecological events may be remembered for long periods of time. These studies also demonstrate the adaptive nature of horses with regard to their ability to generalize over several different testing conditions.
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Rumbaugh, D. M., Riesen, A. H., & Wright, S. C. (1972). Creative responsiveness to objects: a report of a pilot study with young apes. Folia Primatol (Basel), 17(5), 397–403.
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Gaunet, F. (2010). How do guide dogs and pet dogs (Canis familiaris) ask their owners for their toy and for playing? Anim. Cogn., 13(2), 311–323.
Abstract: Abstract When apes are not fully understood by humans, they persist with attempts to communicate, elaborating their behaviours to better convey their meaning. Such abilities have never been investigated in dogs. The present study aimed to clarify any effect of the visual attentional state of the owner on dogs’ (Canis familiaris) social-communicative signals for interacting with humans, and to determine whether dogs persist and elaborate their behaviour in the face of failure to communicate a request. Gaze at a hidden target or at the owner, gaze alternation between a hidden target and the owner, vocalisations and contacts in 12 guide and 12 pet dogs were analysed (i) when the dogs were asked by their owners (blind or sighted) to fetch their inaccessible toy and (ii) when the dogs were subsequently given an unfamiliar object (apparent unsuccessful communication) or their toy (apparent successful communication). No group differences were found, indicating no effect of the visual status of the owner on the dogs’ socio-communicative modes (i.e. no sensitivity to human visual attention). Results, however, suggest that the dogs exhibited persistence (but not elaboration) in their “showing” behaviours in each condition, except that in which the toy was returned. Thus, their communication was about a specific item in space (the toy). The results suggest that dogs possess partially intentional non-verbal deictic abilities: (i) to get their inaccessible toy, the dogs gazed at their owners as if to trigger their attention; gaze alternation between the owner and the target direction, and two behaviours directed at the target were performed, apparently to indicate the location of the hidden toy; (ii) after the delivery of the toy, the dogs behaved as if they returned to the play routine, gazing at their owner whilst holding their toy. In conclusion, this study shows that dogs possess partially intentional non-verbal deictic abilities: they exhibit successive visual orienting between a partner and objects, apparent attention-getting behaviours, no sensitivity to the visual status of humans for communication, and persistence in (but no elaboration of) communicative behaviours when apparent attempts to “manipulate” the human partner fail.
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