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Marr, I., Stefanski, V., & Krueger, K. (2022). Lateralität – ein Indikator für das Tierwohl?[Laterality – an animal welfare indicator?]. Der Praktische Tierarzt, 103(12/2022), 1246–12757.
Abstract: Ein gutes Tierwohl definiert sich nicht nur durch die Abwesenheit von Stressindikatoren, sondern auch durch das Vorhandensein von Indikatoren, die auf ein gutes Wohlergehen hinweisen. So können stressbedingte Erkrankungen vermieden werden. Zur Bestimmung des Tierwohls bei Pferden wurde daher untersucht, inwieweit sich die sensorische Lateralität (einseitiger Gebrauch von Sinnesorganen) und die motorische Lateralität (einseitiger Gebrauch von Gliedmaßen) als einfach, schnell und kostengünstig zu erhebende Parameter eignen. Hierzu werden neben aktueller Literatur auch die eigenen Untersuchungsergebnisse zusammenfassend dargestellt. Die nach außen sichtbar werdende sensorische und motorische Lateralität sind das Resultat der cerebralen Lateralisierung. Dies beinhaltet nicht nur die Aufgabenteilung beider Gehirnhälften für ein effizienteres Aufnehmen und Speichern von Informationen, sondern sie steht auch in Verbindung mit der Entstehung und Verarbeitung von Emotionen, die maßgeblich am Wohlergehen eines Lebewesens beteiligt sind. Kurzzeitige Stressoren führen zu einer Erregung, die je nach Erfahrungen mit positiven oder negativen Emotionen in Verbindung steht. Emotionen helfen dem Organismus dabei, zu überleben. Andauernde negative Emotionen durch regelmäßige oder anhaltende negative Ereignisse führen zu Stress und reduzieren die Erwartung positiver Ereignisse (negativer cognitive Bias). Das Tier ist im Wohlergehen beeinträchtigt. Jüngst zeigte insbesondere die Messung der motorischen Lateralität Potenzial als Indikator für lang anhaltenden und chronischen Stress, denn gestresste Pferde, deren Stresshormonlevel stark ansteigt, zeigen einen zunehmenden Gebrauch der linken Gliedmaßen über einen längeren Zeitraum. Weiterhin zeigen erste Messungen einen Zusammenhang zwischen einer linksseitigen motorischen Lateralität und einer reduzierten Erwartung positiver Ereignisse (negativer cognitive Bias). Zusammen mit der sensorischen Lateralität, die in einer akuten Stressphase ebenso eine Linksverschiebung zeigt und somit als Indikator für Kurzzeitstress gilt, kann eine generelle, vermehrte Linksseitigkeit auch einen Hinweis auf erhöhte Emotionalität und Stressanfälligkeit sein. Eine sich steigernde Linksseitigkeit bedeutet eine präferierte Informationsverarbeitung durch die rechte Gehirnhälfte, die beispielsweise reaktives Verhalten, starke Emotionen und Stressantworten steuert. Es stellte sich jedoch heraus, dass wie bei allen Stressindikatoren auch in der Lateralitätsmessung ein Vergleichswert aus einer vorangegangenen Messung notwendig ist, denn nur Veränderungen zum häufiger werdenden Gebrauch der linken Seite können auf Stress bei Pferden hindeuten und die parallele Erhebung weiterer Parameter, wie zum Beispiel das Verhalten oder Stresshormone, können die Aussage der Lateralität bekräftigen.
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Boissy, A. (1995). Fear and Fearfulness in Animals. The Quarterly Review of Biology, 70(2), 165–191.
Abstract: Persistence of individual differences in animal behavior in reactions to various environmental challenges could reflect basic divergences in temperament, which might be used to predict details of adaptive response. Although studies have been carried out on fear and anxiety in various species, including laboratory, domestic and wild animals, no consistent definition of fearfulness as a basic trait of temperament has emerged. After a classification of the events that may produce a state of fear, this article describes the great variability in behavior and in physiological patterns generally associated with emotional reactivity. The difficulties of proposing fearfulness-the general capacity to react to a variety of potentially threatening situations-as a valid basic internal variable are then discussed. Although there are many studies showing covariation among the psychobiological responses to different environmental challenges, other studies find no such correlations and raise doubts about the interpretation of fearfulness as a basic personality trait. After a critical assessment of methodologies used in fear and anxiety studies, it is suggested that discrepancies among results are mainly due to the modulation of emotional responses in animals, which depend on numerous genetic and epigenetic factors. It is difficult to compare results obtained by different methods from animals reared under various conditions and with different genetic origins. The concept of fearfulness as an inner trait is best supported by two kinds of investigations. First, an experimental approach combining ethology and experimental psychology produces undeniable indicators of emotional reactivity. Second, genetic lines selected for psychobiological traits prove useful in establishing between behavioral and neuroendocrine aspects of emotional reactivity. It is suggested that fearfulness could be considered a basic feature of the temperament of each individual, one that predisposes it to respond similarly to a variety of potentially alarming challenges, but is nevertheless continually modulated during development by the interaction of genetic traits of reactivity with environmental factors, particularly in the juvenile period. Such interaction may explain much of the interindividual variability observed in adaptive responses.
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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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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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von Borstel, U. U., Duncan, I. J. H., Shoveller, A. K., Merkies, K., Keeling, L. J., & Millman, S. T. (2009). Impact of riding in a coercively obtained Rollkur posture on welfare and fear of performance horses. Appl. Anim. Behav. Sci., 116(2-4), 228–236.
Abstract: Rollkur, the usually coercively obtained hyperflexion of the horse's neck, is employed as a training method by some dressage riders; however, its use is controversial as it may cause discomfort and adversely affect the horse's welfare. The objectives of this study were to determine: (1) if horses showed differences in stress, discomfort and fear responses as measured by heart rate and behaviour when ridden in Rollkur (R) obtained by pressure on the reins compared to regular poll flexion (i.e. with the nose-line being at or just in front of the vertical; N), and (2) if they showed a preference between the two riding styles when given the choice. Fifteen riding horses were ridden 30 times through a Y-maze randomly alternating between sides. Riding through one arm of the Y-maze was always followed by a short round ridden in R, whereas riding through the other arm was followed by a short round ridden in N. Immediately after the conditioning phase, horses were again repeatedly ridden into the maze; however, riders left it to the horse to decide which arm of the maze to enter. During R, horses moved slower and showed more often behavioural signs of discomfort, such as tail-swishing, head-tossing or attempted bucks (P < 0.05), and 14 of the 15 horses chose significantly (P < 0.05) more often the maze-arm associated with N rather than R. Subsequently, eight of the horses were also subjected to two fear tests following a short ride in N as well as a ride in R. During R, horses tended to react stronger (P = 0.092) to the fear stimuli and to take longer (P = 0.087) to approach them. These findings indicate that a coercively obtained Rollkur position may be uncomfortable for horses and that it makes them more fearful and therefore potentially more dangerous to ride. Further studies need to assess horses' reaction to gradual training of Rollkur, as opposed to a coercively obtained hyperflexion, in order to decide whether the practice should be banned.
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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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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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Carson, K., & Wood-Gush, D. G. M. (1983). Equine behaviour: I. A review of the literature on social and dam--Foal behaviour. Applied Animal Ethology, 10(3), 165–178.
Abstract: In most cases, the social organisation of each of the seven species of Equidae existing today outside captivity is either territorial or non-territorial. The striking differences found between these two types of organisation in the social grouping and bonds, mating behaviour, leadership and dominance hierarchies of the animals are examined. It is thought that the non-territorial species show a less primitive type of organisation than the territorial animals. Infant Equidae are precocious animals and are able to follow their dams soon after birth. They stay close by their dams and travel with the herd from an early age and are therefore classified as “followers”, in contrast to the species which have a period of hiding after birth. Dams recognise their foals immediately after birth, whereas it takes 2 or 3 days for a foal to form an attachment to its dam. Being in close proximity to their dams, foals are able to nurse frequently and, unless artificially weaned, a foal will nurse until its dam foals again. Foals start to graze during their first week and as they grow older they spend more time grazing and less time nursing and resting. It is normal for foals to be corprophagic until one month old, and this provides them with bacteria essential for the digestion of fibre. Play behaviour is solitary in very young foals, but after 4 weeks of age, foals play together, with male foals playing more than females and showing more aggressive, fighting movements in play.
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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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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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