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Christensen, J. W., Munk, R., Hawson, L., Palme, R., Larsen, T., Egenvall, A., et al. (2021). Rider effects on horses' conflict behaviour, rein tension, physiological measures and rideability scores. Appl. Anim. Behav. Sci., 234, 105184.
Abstract: Many breeding organisations include a subjective scoring of rideability by a professional rider into their evaluation of sports horses, but the consistency and reliability of the scoring system is debateable. The aim of this study was to investigate (i) whether professional riders agree in their scoring of rideability, and (ii) whether rideability scores are affected by rein tension, horse conflict behaviour, heart rate, and salivary cortisol, and (iii) whether riders induce different levels of conflict behaviour and physiological responses in the horses. Ten professional, female riders each rode 10 dressage horses (level M German scale; n = 100 combinations) through a standardised dressage test (10 min warm-up followed by a 4-min test) and subsequently scored the horses for rideability on the official 1-10 scale (1 = poor to 10 = excellent) from the Danish Riding Federation. Rein tension, horse heart rate, saliva cortisol and conflict behaviour were measured for each rider-horse pair. The riders were inconsistent in their scoring of rideability to the individual horses, e.g. scores for one of the horses ranged from 1 to 8. There was a significant effect of rider (P = 0.003) and the frequency of conflict behaviour (undesired head movements: P < 0.001, breaking the gait: P = 0.013, and other evasive behaviour: P = 0.032) on rideability scores, i.e. the more conflict behaviour the lower the score. There was no significant effect of rein tension and the physiological measures on rideability scores. However, there was a significant effect of rider on rein tension, horses' heart rate and increases in saliva cortisol concentrations and a tendency for some types of conflict behaviour, suggesting that some riders induced more discomfort in the horses. Future studies could help shed light on which elements of riding style are particularly important for sports horse welfare. In conclusion, this study found a large variation in rideability scores assigned to ten sports horses by ten professional riders. Rideability scores were dependent on the level of horse conflict behaviour, but not rein tension and physiological measures. Further studies are needed to improve the objectivity, consistency and reliability of rideability assessment of sports horses.
Keywords: Equitation science; Heart rate; Horse riding; Sport horse; Temperament; Training cues
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da Cruz, A. B., Hirata, S., dos Santos, M. E., & Mendonça, R. S. (2023). Show me your best side: Lateralization of social and resting behaviors in feral horses. Behav. Process., 206, 104839.
Abstract: Growing evidence shows a variety of sensorial and motor asymmetries in social and non-social interactions in various species, indicating a lateralized processing of information by the brain. Using digital video cameras on tripods and drones, this study investigated lateralization in frequency and duration of social behavior patterns, in affiliative, agonistic, and resting contexts, in a feral population of horses (Equus ferus caballus) in Northern Portugal, consisting of 37 individuals organized in eight harem groups. Affiliative interactions (including grooming) were more often performed, and lasted longer, when recipients were positioned to the right side. In recumbent resting (animals lying down) episodes on the left side lasted longer. Our results of an affiliative behavior having a right side tendency, provide partial support to the valence-specific hypothesis of Ahern and Schwartz (1979) – left hemisphere dominance for positive affect, affiliative behaviors. Longer recumbent resting episodes on the left side may be due to synchronization. However, in both instances it is discussed how lateralization may be context dependent. Investigating the position asymmetries of social behaviors in feral equids will contribute to a better understanding of differential lateralization and hemispheric specialization from the ecological and evolutionary perspectives.
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Goursot, C., Düpjan, S., Puppe, B., & Leliveld, L. M. C. (2021). Affective styles and emotional lateralization: A promising framework for animal welfare research. Appl. Anim. Behav. Sci., 237, 105279.
Abstract: The growing recognition of animals as individuals has broader implications for farm animal welfare research. Even under highly standardized on-farm conditions, farm animals show heterogeneous but individually consistent behavioural patterns towards various stimuli, based on how they appraise these stimuli. As a result, animal welfare is likely to be highly individual as well, and studying the proximate mechanisms underlying distinct individual behaviour patterns and appraisal will improve animal welfare research. We propose to extend the framework of affective styles to bridge the gap between existing research fields on animal personality and affective states. Affective styles refer to consistent individual differences in emotional reactivity and regulation and can be predicted by baseline cerebral lateralization. Likewise, animals with consistent left or right motor biases--a proxy measure of individual patterns in cerebral lateralization--have been shown to differ in their personality, emotional reactivity, motivational tendencies or coping styles. In this paper, we present the current knowledge of the links between laterality and stable individual traits in behaviour and affect in light of hypotheses on emotional lateralization. Within our suggested framework, we make recommendations on how to investigate affective styles in non-human animals and give practical examples. This approach has the potential to promote a science of affective styles in nonhuman animals and significantly advance research on animal welfare.
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Leliveld, L. M. C., Düpjan, S., Tuchscherer, A., & Puppe, B. (2020). Hemispheric Specialization for Processing the Communicative and Emotional Content of Vocal Communication in a Social Mammal, the Domestic Pig. Front. Behav. Neurosci., 14, 596758.
Abstract: In humans, speech perception is lateralized, with the left hemisphere of the brain dominant in processing the communicative content and the right hemisphere dominant in processing the emotional content. However, still little is known about such a division of tasks in other species. We therefore investigated lateralized processing of communicative and emotionally relevant calls in a social mammal, the pig (Sus scrofa). Based on the contralateral connection between ears and hemispheres, we compared the behavioural and cardiac responses of 36 young male pigs during binaural and monaural (left or right) playback to the same sounds. The playback stimuli were calls of social isolation and physical restraint, whose communicative and emotional relevance, respectively, were validated prior to the test by acoustic analyses and during binaural playbacks. There were indications of lateralized processing mainly in the initial detection (left head-turn bias, indicating right hemispheric dominance) of the more emotionally relevant restraint calls. Conversely, there were indications of lateralized processing only in the appraisal (increased attention during playback to the right ear) of the more communicative relevant isolation calls. This implies differential involvement of the hemispheres in the auditory processing of vocalizations in pigs and thereby hints at similarities in the auditory processing of vocal communication in non-human animals and speech in humans. Therefore, these findings provide interesting new insight in the evolution of human language and auditory lateralization.
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Camerlink, I., Menneson, S., Turner, S. P., Farish, M., & Arnott, G. (2018). Lateralization influences contest behaviour in domestic pigs. Scientific Reports, 8(1), 12116.
Abstract: Cerebral lateralization, i.e. hemispheric asymmetries in structure and function, relates in many species to a preference to attack from their left. Lateralization increases cognitive capacity, enabling the simultaneous processing of multiple sources of information. Therefore, lateralization may constitute a component of fighting ability (Resource Holding Potential), and/or influence the efficiency of information-gathering during a contest. We hypothesized that lateralization will affect contest outcome and duration, with an advantage for more strongly lateralized individuals. In 52 dyadic contests between weight-matched pigs (Sus scrofa; n = 104; 10 wk age), the direction of orientation towards the opponent was scan sampled every 10 s. Laterality indexes (LI) were calculated for the direction and strength of lateralization. Up to 12.5% of the individuals showed significant lateralization towards either the right or left but lateralization was absent at the population level. In line with our hypothesis, animals showing strong lateralization (irrespective of direction) had a shorter contest duration than animals showing weak lateralization. Winners did not differ from losers in their strength or direction of lateralization. Overall the results suggest that cerebral lateralization may aid in conflict resolution, but does not directly contribute to fighting ability, and will be of value in the study of animal contests.
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Versace, E., Morgante, M., Pulina, G., & Vallortigara, G. (2007). Behavioural lateralization in sheep (Ovis aries). Behav. Brain. Res., 184(1), 72–80.
Abstract: This study investigates behavioural lateralization in sheep and lambs of different ages. A flock was tested in a task in which the animals were facing an obstacle and should avoid it on either the right or left side to rejoin flock-mates (adult sheep) or their mothers (lambs). A bias for avoiding the obstacle on the right side was observed, with lambs apparently being more lateralized than sheep. This right bias was tentatively associated with the left-hemifield laterality in familiar faces recognition which has been documented in this species. Differences between adult sheep and lambs were likely to be due to differences in social reinstatement motivation elicited by different stimuli (flock-mates or mothers) at different ages. Preferential use of the forelegs to step on a wood-board and direction of jaw movement during rumination was also tested in adult animals. No population bias nor individual-level lateralization was observed for use of the forelegs. At the same time, however, there was a large number of animals showing individual-level lateralization for the direction of jaw movement during rumination even though there was no population bias. These findings highlight that within the same species individual- and population-level lateralization can be observed in different tasks. Moreover, the results fit the general hypothesis that population-level asymmetries are more likely to occur in tasks that require social coordination among behaviourally asymmetric individuals.
Keywords: Lateralization; Laterality; Brain asymmetry; Hemisphere; Sheep; Lamb; Strength of lateralization
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Meek, P. D., Ballard, G. - A., & Fleming, P. J. S. (2015). The pitfalls of wildlife camera trapping as a survey tool in Australia. Aust. Mammal., 37(1), 13–22.
Abstract: Camera trapping is a relatively new addition to the wildlife survey repertoire in Australia. Its rapid adoption has been unparalleled in ecological science, but objective evaluation of camera traps and their application has not kept pace. With the aim of motivating practitioners to think more about selection and deployment of camera trap models in relation to research goals, we reviewed Australian camera trapping studies to determine how camera traps have been used and how their technological constraints may have affected reported results and conclusions. In the 54 camera trapping articles published between 1991 and 2013, mammals (86%) were studied more than birds (10%) and reptiles (3%), with small to medium-sized mammals being most studied. Australian camera trapping studies, like those elsewhere, have changed from more qualitative to more complex quantitative investigations. However, we found that camera trap constraints and limitations were rarely acknowledged, and we identified eight key issues requiring consideration and further research. These are: camera model, camera detection system, camera placement and orientation, triggering and recovery, camera trap settings, temperature differentials, species identification and behavioural responses of the animals to the cameras. In particular, alterations to animal behaviour by camera traps potentially have enormous influence on data quality, reliability and interpretation. The key issues were not considered in most Australian camera trap papers and require further study to better understand the factors that influence the analysis and interpretation of camera trap data and improve experimental design.
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Dyson, S., Berger, J., Ellis, A. D., & Mullard, J. (2018). Development of an ethogram for a pain scoring system in ridden horses and its application to determine the presence of musculoskeletal pain. Journal of Veterinary Behavior, 23, 47–57.
Abstract: There is evidence that more than 47% of the sports horse population in normal work may be lame, but the lameness is not recognized by owners or trainers. An alternative means of detecting pain may be recognition of behavioral changes in ridden horses. It has been demonstrated that there are differences in facial expressions in nonlame and lame horses. The purpose of this study was to develop a whole horse ethogram for ridden horses and to determine whether it could be applied repeatedly by 1 observer (repeatability study, 9 horses) and if, by application of a related pain behavior score, lame horses (n = 24) and nonlame horses (n = 13) could be differentiated. It was hypothesized that there would be some overlap in pain behavior scores among nonlame and lame horses; and that overall, nonlame horses would have a lower pain behavior score than lame horses. The ethogram was developed with 117 behavioral markers, and the horses were graded twice in random order by a trained specialist using video footage. Overall, there was a good correlation between the 2 assessments (P < 0.001; R2 = 0.91). Behavioral markers that were not consistent across the 2 assessments were omitted, reducing the ethogram to 70 markers. The modified ethogram was applied to video recordings of the nonlame horses and lame horses (ethogram evaluation). There was a strong correlation between 20 behavioral markers and the presence of lameness. The ethogram was subsequently simplified to 24 behavioral markers, by the amalgamation of similar behaviors which scored similarly and by omission of markers which showed unreliable results in relation to lameness. Following this, the maximum individual occurrence score for lame horses was 14 (out of 24 possible markers), with a median and mean score of 9 (±2 standard deviation) compared with a maximum score of 6 for nonlame horses, with a median and mean score of 2 (±1.4). For lame horses, the following behaviors occurred significantly more (P < 0.05, chi-square): ears back, mouth opening, tongue out, change in eye posture and expression, going above the bit, head tossing, tilting the head, unwillingness to go, crookedness, hurrying, changing gait spontaneously, poor quality canter, resisting, and stumbling and toe dragging. Recognition of these features as potential indicators of musculoskeletal pain may enable earlier recognition of lameness and avoidance of punishment-based training. Further research is necessary to verify this new ethogram for assessment of pain in ridden horses.
Keywords: Lameness; Equine behavior; Pain grading; Headshaking; Bucking; Rearing
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Bussières, G., Jacques, C., Lainay, O., Beauchamp, G., Leblond, A., Cadoré, J. - L., et al. (2008). Development of a composite orthopaedic pain scale in horses. Research in Veterinary Science, 85(2), 294–306.
Abstract: This study addresses development and validation of a composite multifactorial pain scale (CPS) in an experimental equine model of acute orthopaedic pain. Eighteen horses were allocated to control (sedation with/without epidural analgesia – mixture of morphine, ropivacaine, detomidine and ketamine) and experimental groups: amphotericin-B injection in the tarsocrural joint induced pain and analgesia was either i.v. phenylbutazone administered post-induction of synovitis, or pre-emptive epidural mixture, or a pre-emptive combination of the 2. Inter- and intra-observer reproducibility was good (0.8<K<1). The key specific and sensitive behavioural indices were response to palpation of the painful area, posture, and, of lesser value, pawing on the floor, kicking at abdomen and head movement. Of particular interest was the statistical correlation observed between the CPS and both non-invasive blood pressure (P<0.0001) and blood cortisol (P<0.002). This study established the value of some behavioural and physiological criteria in determining equine orthopaedic pain intensity and clearly demonstrated that pre-emptive, multimodal analgesia provided better management than the two other protocols tested.
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Krueger, K., Gruentjens, T., & Hempel, E. (2023). Wolf contact in horses at permanent pasture in Germany. Plos One, 18(8), e0289767.
Abstract: Wolves returned to Germany in 2000, leading to fear in German horse owners that their horses could be in danger of wolf attacks or panic-like escapes from pastures when sighting wolves. However, reports from southern European countries indicate that wolf predation on horses diminishes with increasing presence of wildlife. Therefore, we conducted a long-term, filed observation between January 2015 and July 2022 on 13 non breeding riding horses, mares and geldings, kept permanently on two pastures within the range of wildlife and a stable wolf pack with annual offspring. Wildlife cameras at the fences of the pastures made 984 times recordings of wolves and 3151 times recordings of wildlife in and around the pastures. Between 1 January 2022 and 23 March 2022 we observed two stable horse groups. Pasture 1 was grazed by five horses of mixed breed, four mares and one gelding, with the median age of 8 years (min. = 6y, max. = 29y). Pasture 2 was grazed by eight heavy warmbloods and draught horses, three mares and five geldings, with the median age of 16 years (min. = 13y, max. = 22y). During this period no wolf was recorded at pasture 2, but wild boar several times, whereas at pasture 1, wolves were recorded 89 times, and for the wildlife mostly hare. Wolves may have avoided pasture 2 because of the presence of wild boar or because the large group of older, heavy breed horses may have formed a stable, protective group. The latter needs to be confirmed in a follow-up field observation, which records anti-predator behavior and welfare indicators in horses. In conclusion, wolves did not attack the mature horses on pastures with plenty of wildlife and the horses did not respond to the presence of wolves with visible signs of reduced welfare or panic. This indicates that wolves may prefer to prey on easily accessible wildlife around and at horse pastures and that Central European horses become accustom to the presence of non-hunting wolves.
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