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Christensen, J. W., & Rundgren, M. (2008). Predator odour per se does not frighten domestic horses. Appl. Anim. Behav. Sci., 112(1-2), 136–145.
Abstract: Horses frequently react nervously when passing animal production farms and other places with distinctive smells, leading riders to believe that horses are innately frightened by certain odours. In three experiments, we investigated how horses respond to (1) urine from wolves and lions, (2) blood from slaughtered conspecifics and fur-derived wolf odour, and (3) a sudden auditory stimulus in either presence or absence of fur-derived wolf odour. The experiments were carried out under standardised conditions using a total of 45 naive, 2-year-old horses. In the first two experiments we found that horses showed significant changes in behaviour (Experiments 1 and 2: increased sniffing; Experiment 2 only: increased vigilance, decreased eating, and more behavioural shifts), but no increase in heart rate compared to controls when exposed to predator odours and conspecific blood in a known test environment. However, the third experiment showed that exposure to a combination of wolf odour and a sudden stimulus (sound of a moving plastic bag) caused significantly increased heart rate responses and a tendency to a longer latency to resume feeding, compared to control horses exposed to the sudden stimulus without the wolf odour. The results indicate that predator odour per se does not frighten horses but it may cause an increased level of vigilance. The presence of predator odour may, however, cause an increased heart rate response if horses are presented to an additional fear-eliciting stimulus. This strategy may be adaptive in the wild where equids share habitats with their predators, and have to trade-off time and energy spent on anti-predation responses against time allocated to essential non-defensive activities.
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Christensen, J. W., Søndergaard, E., Thodberg, K., & Halekoh, U. (2011). Effects of repeated regrouping on horse behaviour and injuries. Appl. Anim. Behav. Sci., 133(3), 199–206.
Abstract: Domestic horses are faced with social challenges throughout their lives due to limitations in social contact, space restrictions and frequent changes in social companionship. This is in contrast to natural conditions where horses live in relatively stable harem bands. Currently, little is known about how repeated regrouping affect horse behaviour and welfare, and it is unknown whether horses may adapt to regrouping. In this study, we aimed to investigate the effects of an unstable group structure, caused by weekly regroupings, on behaviour and frequency of injuries in young horses. Forty-five horses were included in the study and were randomly assigned to the treatments; Stable (S; seven groups of three horses) or Unstable (U; eight groups of three horses). The experimental period lasted 7 weeks, during which horses in Stable groups remained in the same group, whereas one horse was exchanged between Unstable groups every week. The groups were kept in 80m×80m grass-covered enclosures and were fed additional roughage on the ground daily. Social interactions were recorded in Unstable groups immediately after each regrouping (30min), and in both Stable and Unstable groups on day 1, 3 and 6 after each regrouping (2×20min/group/day). Injuries were scored by the end of the experimental period. The level of aggression shown by horses in Unstable groups immediately after regrouping was not affected by week (F5,35=0.42, P=0.83), indicating that horses neither habituated, nor sensitized, to repeated regrouping. Compared to horses in Stable groups, more agonistic behaviour was shown by horses in Unstable groups (i.e. non-contact agonistic; F1,65=5.60, P=0.02), whereas there was no treatment effect on other variables. The level of play behaviour appeared, however, to be more variable in Unstable groups. There was a significant effect of week on the level of contact agonistic interactions as well as greeting behaviour, due to a high occurrence in weeks 4-6. Non-contact agonistic interactions constituted the major part of agonistic interactions (66%). Possibly as consequence, no serious injuries were registered and there was no treatment effect (U=184; P=0.11). We conclude that the behaviour of young horses is affected by group management, and that horses appear not to adapt to weekly regroupings.
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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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Christensen, J. W., Zharkikh, T., & Ladewig, J. (2008). Do horses generalise between objects during habituation? Appl. Anim. Behav. Sci., 114(3-4), 509–520.
Abstract: Habituation to frightening stimuli plays an important role in horse training. To investigate the extent to which horses generalise between different visual objects, 2-year-old stallions were habituated to feeding from a container placed inside a test arena and assigned as TEST (n=12) or REFERENCE horses (n=12). In Experiment 1, TEST horses were habituated to six objects (ball, barrel, board, box, cone, cylinder) presented in sequence in a balanced order. The objects were of similar size but different colour. Each object was placed 0.5m in front of the feed container, forcing the horses to pass the object to get to the food. TEST horses received as many 2min exposures to each object as required to meet a habituation criterion. We recorded behavioural reactions to the object, latency to feed, total eating time, and heart rate (HR) during all exposures. There was no significant decrease in initial responses towards a novel object with increasing object number, indicating that habituation was stimulus-specific. REFERENCE horses were exposed to the test arena without objects in the same period, and reactions of TEST and REFERENCE horses were subsequently compared in a fear-test (2m2.5m rubber mat placed under the feed container, forcing the horses to step on the mat to get food). There were no significant differences between the treatment groups, i.e. previous habituation of TEST horses to six visual objects did not reduce responses in a fear-test involving visual and tactile stimulation. Due to the lack of generalisation in Experiment 1, we designed a supplemental experiment (Experiment 2), in which REFERENCE horses were exposed to the same six objects except that object colour was kept constant. We found a significant reduction in response (behaviour and HR) with increasing object number, indicating that horses generalise between similarly coloured objects of varying shape. We conclude that a high degree of object similarity, e.g. identical colouring, appears to be crucial for object generalisation in horses.
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Christensen, J. W., Zharkikh, T., Ladewig, J., & Yasinetskaya, N. (2002). Social behaviour in stallion groups (Equus przewalskii and Equus caballus) kept under natural and domestic conditions. Appl. Anim. Behav. Sci., 76(1), 11–20.
Abstract: The aim of this study was to investigate social behaviour in differently reared stallions in their respective environments; one group of stallions was reared under typical domestic conditions whereas the other group was reared and lives under natural conditions. The domestic group consisted of 19, 2-year-old stallions (Equus caballus), which were all weaned at 4 months of age and experienced either individual or group housing facilities before being pastured with the other similarly aged stallions. The natural living and mixed age group of Przewalski stallions (E. przewalskii) consisted of 13 stallions, most of which were juveniles (n=11, <=4 years; n=2, >9 years). The domestic group was studied in a 4-ha enclosure at the Danish Institute of Agricultural Sciences and the Przewalski group under free-ranging conditions in a 75-ha enclosure in the Askania Nova Biosphere Reserve, Ukraine. Behavioural data was collected during 168 h of direct observation. The occurrence of 14 types of social interactions was recorded and group spacing behaviour was studied using nearest neighbour recordings. In spite of very different environments, reflecting domestic and natural rearing conditions, many similarities in behaviour was found. Play and play fight behaviour was very similar in the two stallion groups. Quantitative differences were found in social grooming since Przewalski stallions groomed more frequently (P=0.004), and in investigative behaviours, since domestic stallions showed more nasal (P=0.005) and body sniffing (P<0.001), whereas Przewalski stallions directed more sniffing towards the genital region (P<0.001). These differences may, however, be attributed to environmental factors and in the period of time the stallions were together prior to the study period. Quantitative differences appeared in some agonistic behaviours (kick threat, P<0.001; and kick, P<0.001), but data do not support earlier findings of Przewalski horses being significantly more aggressive than domestic horses. In general, Przewalski stallions engaged in more social interactions, and they showed less group spacing, i.e. maintained a significantly shorter distance between neighbours (P<0.001). The study indicates that also domestic horses, which have been reared under typical domestic conditions and allowed a period on pasture, show social behaviour, which is very similar to that shown by their non-domestic relatives.
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Church, D. L., & Plowright, C. M. S. (2006). Spatial encoding by bumblebees (Bombus impatiens) of a reward within an artificial flower array. Anim. Cogn., 9(2), 131–140.
Abstract: We presented bumblebees a spatial memory task similar to that used with other species (e.g., cats, dogs, and pigeons). In some conditions we allowed for presence of scent marks in addition to placing local and global spatial cues in conflict. Bumblebees (Bombus impatiens) were presented an array of artificial flowers within a flight cage, one flower offering reward (S+), while the others were empty (S-). Bees were tested with empty flowers. In Experiment 1, flowers were either moved at the time of testing or not. Bees returned to the flower in the same absolute position of the S+ (the flower-array-independent (FAI) position), even if it was in the wrong position relative to the S- and even when new flower covers prevented the use of possible scent marks. New flower covers (i.e., without possible scent marks) had the effect of lowering the frequency of probing behavior. In Experiment 2, the colony was moved between training and testing. Again, bees chose the flower in the FAI position of the S+, and not the flower that would be chosen using strictly memory for a flight vector. Together, these experiments show that to locate the S+ bees did not rely on scent marks nor the positions of the S-, though the S- were prominent objects close to the goal. Also, bees selected environmental features to remember the position of the S+ instead of relying upon a purely egocentric point of view. Similarities with honeybees and vertebrates are discussed, as well as possible encoding mechanisms.
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Church, R. M. (1997). Quantitative models of animal learning and cognition. J Exp Psychol Anim Behav Process, 23(4), 379–389.
Abstract: This article reviews the prerequisites for quantitative models of animal learning and cognition, describes the types of models, provides a rationale for the development of such quantitative models, describes criteria for their evaluation, and makes recommendations for the next generation of quantitative models. A modular approach to the development of models is described in which a procedure is considered as a generator of stimuli and a model is considered as a generator of responses. The goal is to develop models that, in combination with many different procedures, produce sequences of times of occurrence of events (stimuli and responses) that are indistinguishable from those produced by the animal under many experimental procedures and data analysis techniques.
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Cinková, I., & Policht, R. (2016). Sex and species recognition by wild male southern white rhinoceros using contact pant calls. Anim. Cogn., 19(2), 375–386.
Abstract: Recognition of information from acoustic signals is crucial in many animals, and individuals are under selection pressure to discriminate between the signals of conspecifics and heterospecifics or males and females. Here, we first report that rhinos use information encoded in their calls to assess conspecifics and individuals of closely related species. The southern (Ceratotherium simum) and critically endangered northern (C. cottoni) white rhinos are the most social out of all the rhinoceros species and use a contact call pant. We found that southern white rhino pant calls provide reliable information about the caller’s sex, age class and social situation. Playback experiments on wild territorial southern white rhinoceros males revealed that they responded more strongly to the pant calls of conspecific females compared to the calls of other territorial males. This suggests that pant calls are more important form of communication between males and females than between territorial males. Territorial southern males also discriminated between female and territorial male calls of northern species and reacted more intensively to the calls of northern than southern males. This might be caused by a novelty effect since both species naturally live in allopatry. We conclude that white rhinos can directly benefit from assessing individuals at long distances using vocal cues especially because their eyesight is poor. Pant calls thus likely play a significant role in their social relationships and spatial organization. In addition, better understanding of vocal communication in white rhinos might be helpful in conservation management particularly because of their low reproduction in captivity.
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Cinková, I., & Policht, R. (2015). Discrimination of familiarity and sex from chemical cues in the dung by wild southern white rhinoceros. Anim. Cogn., 18(1), 385–392.
Abstract: Communication in rhinos is primarily mediated by the vocal and olfactory signals as they have relatively poor eyesight. White rhinos are the most social of all the rhinoceros species, they defecate at common dungheaps and the adult bulls use dung and urine to mark their territory. Chemical communication may therefore be particularly important in the social interactions of white rhinos, and its knowledge could be very helpful in their management and conservation. However, no studies have investigated up until now the olfactory discrimination in any rhinoceros species in the wild. We have experimentally studied the reactions of the wild southern white rhinos (Ceratotherium simum) to the dung of familiar and unfamiliar adult females and adult territorial males. We registered the number of sniffing events, the duration of sniffing and the latency of the vigilance posture from the onset of sniffing. The dung of unfamiliar rhinos was sniffed longer than that of familiar rhinos. The rhinos showed a shorter latency of vigilance posture to the familiar dung of males than that of females. For unfamiliar dung, they displayed a shorter latency of vigilance posture to female than male dung. Our results indicate that the rhinos are able to discriminate the familiarity and sex of conspecifics from the smell of their dung. Olfactory cues could therefore play an important role in the social relationships and spatial organization of the southern white rhinoceros.
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Clara, E., Regolin, L., Vallortigara, G., & Rogers, L. (2007). Perception of the stereokinetic illusion by the common marmoset (Callithrix jacchus). Anim. Cogn., 10(2), 135–140.
Abstract: Stereokinetic illusions have never been investigated in non-human primates, nor in other mammalian species. These illusions consist in the perception of a 3D solid object when certain 2D stimuli are rotated slowly in the plane perpendicular to the line of sight. The ability to perceive the stereokinetic illusion was investigated in the common marmoset (Callithrix jacchus). Four adult marmosets were trained to discriminate between a solid cylinder and a solid cone for food reward. Once learning criterion was reached, the marmosets were tested in sets of eight probe trials in which the two solid objects used at training were replaced by two rotating 2D stimuli. Only one of these stimuli produced, at least to the human observer, the stereokinetic illusion corresponding to the solid object previously reinforced. At test, the general behaviour and the total time spent by the marmosets observing each stimulus were recorded. The subjects stayed longer near the stimulus producing the stereokinetic illusion corresponding to the solid object reinforced at training than they did near the illusion corresponding to the previously non-rewarded stimulus. Hence, the common marmosets behaved as if they could perceive stereokinetic illusions.
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