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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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Janczarek, I., Wisniewska, A., Chruszczewski, M. H., Tkaczyk, E., & Górecka-Bruzda, A. (2020). Social Behaviour of Horses in Response to Vocalisations of Predators. Animals, 10(2331).
Abstract: We tested the hypothesis that social defensive responses to the vocalisation of a predator still exist in horses. The recordings of a grey wolf, an Arabian leopard and a golden jackal were played to 20 Konik polski and Arabian mares. Durations of grazing, standing still, standing alert and the number of steps in walk and trot/canter were measured. In one-minute scans, the distances of the focal horse from the reference horse (DIST-RH) and from the nearest loudspeaker (DIST-LS) were approximated. The vocalisation of a leopard aroused the Arabians more than the Koniks (less grazing, stand-still and walk, more stand-alert and trotting/cantering). Koniks showed more relaxed behaviours to the leopard vocalisation (more grazing, stand-still and walk), but high alertness to the wolf playback (stand-alert, trotting/cantering). Spatial formation of the herd of Koniks showed tight grouping (lower DIST-RH) and maintaining distance from the potential threat (DIST-LS) in response to the wolf howling, while the Arabians approached the loudspeakers in linear herd formation when the leopard growls were played. Adult horses responded to potential predation by changing spatial group formations. This ability to apply a social strategy may be one of the explanations for the least number of horses among all hunted farm animal species.
Keywords: horse; predator; vocalisation; social defensive behaviour; grey wolf; Arabian leopard; golden jackal
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Niederhöfer, S. (2009). Stressbelastung bei Pferden in Abhängigkeit des Haltungssystems. Ph.D. thesis, Tierärztliche Hochschule Hannover, Hannover.
Abstract: Pferdegerechte Haltungssysteme spielen bei der heutigen Nutzung des Freizeit- oder Sportpartners Pferd eine wichtige Rolle, da naturnahe Haltungsbedingungen nur in seltenen Fällen zu realisieren sind. Sämtliche Aufstallungsformen müssen als Kompromiss angesehen werden und bieten somit Vor- und Nachteile. Die vorliegende Untersuchung verfolgte den Zweck, die Stressbelastung von Pferden in verschiedenen Haltungssystemen zu erfassen und vergleichend zu beurteilen. Während der von Mai 2006 bis Juli 2007 durchgeführten Versuche wurden 24 zwei- bis dreijährige Hannoveranerstuten für jeweils 4 Wochen in insgesamt 6 verschiedenen Haltungsformen aufgestallt. Die Haltungsvarianten beinhalteten die Einzelbox (Variante 1), die Einzelbox mit einem frei zugänglichem Paddock (Variante 2), die gemeinsame Haltung von zwei Pferden in zwei Boxen und einem angeschlossenen frei zugänglichem Auslauf (Variante 3) und die Gruppenhaltung von 6 Pferden in einer Mehrraumauslaufhaltung (Variante 4 – 6). Die Haltungsvariante 4 differierte von den Varianten 5 und 6 durch seinen ungegliederten Liegebereich. In den Variante 5 und 6 wurde der Liegebereich der Gruppenhaltung durch eine über die halbe Breite reichende Trennwand in zwei Bereiche unterteilt. Die Trennwand befand sich in Haltungsvariante 5 im rechten Winkel an die Außenwand grenzend, während sie in Variante 6 an der der Stallgasse zugewandten Seite des Liegebereichs aufgestellt wurde. Um eine Vergleichbarkeit mit einem Reitpferd zu schaffen, wurden alle Pferde in allen Haltungssystemen täglich für circa eine Stunde in einer Freilaufanlage bewegt. Die Stressbelastung wurde mittels der Messung der Herzfrequenzvariabilität (Parameter pNN50 und SD1) und der Bestimmung fäkaler Cortisolmetaboliten erfasst. Zusätzlich wurden Videoauswertungen hinsichtlich der Beobachtung von Verhaltensauffälligkeiten in den Haltungsvarianten 1 und 2 durchgeführt. Die durchschnittlich niedrigste Stressbelastung erfuhren die Pferde durch die Haltungsvarianten 4 (pNN50: 46,32 %, SD1:158,58 ms, Cortisolmetaboliten: 21,01nmol/kg Kot) und 6 (pNN50: 47,1 %, SD1: 144,62 ms, Cortisolmetaboliten: 21,01 nmol/kg Kot). Die
141 Auswertung der pNN50- und Cortisolmetabolitenwerte ergab die größte Stressbelastung der Pferde in den Varianten 1 (pNN50: 42,81 %, SD1: 134,52 ms, Cortisolmetaboliten: 28,56 nmol/kg Kot) und 3 (pNN50: 42,41 %, SD1: 135,36, Cortisolmetaboliten: 28,60 nmol/kg Kot). Die Auswertung der SD1-Werte zeigte zusätzlich noch eine hohe Stressbelastung der Pferde in der Variante 5 (pNN50: 44,83 %, SD1: 119,24 ms, Cortisolmetaboliten: 27,18 nmol/kg Kot). Die Haltungsvariante 2 (pNN50: 45,77 %, SD1: 144,25 ms, Cortisolmetaboliten: 27,59 nmol/kg Kot) beziehungsweise die Varianten 2 und 5 (bei der Betrachtung der pNN50- und der Cortisolmetabolitenwerte) verursachten im Durchschnitt eine mittlere Stressbelastung. Teilweise waren die Unterschiede jedoch zu gering um die Signifikanzgrenze zu überschreiten. Die Betrachtung der einzelnen Pferde und der Pferdegruppen ergab große interindividuelle Unterschiede und deutliche gruppenspezifische Gemeinsamkeiten. Die Position eines Tieres in der Rangordnung hatte in diesen Untersuchungen bei der Betrachtung der Mittelwerte keinen Einfluss auf die Stressbelastung in Abhängigkeit der Haltungsvariante. Vielmehr zeigten sich auch hier deutliche individuelle Unterschiede in der Stressanfälligkeit und den Vorlieben der einzelnen Pferde bezüglich der verschiedenen Haltungssysteme. Die Analyse der Videoaufzeichnungen ergab, dass mehrere Pferde Kreisbewegungen in der Einzelbox ohne Auslauf (Variante 1) zeigten, während in der Variante 2 (Paddockbox) kein Pferd durch Kreisbewegungen auffiel. Andere Verhaltensauffälligkeiten oder gar Verhaltensstörungen wurden nicht beobachtet. Bei der Auswertung der Aufenthaltshäufigkeit und der Aufenthaltesdauer in den 4 Boxenquadranten zeigten sich einige Pferde sehr aktiv, was ein Hinweis auf eventuelle Unruhe oder Nervosität sein kann, während andere Pferde im Vergleich zum Gruppendurchschnitt sehr ruhig wirkten, da sie sich in der Box kaum bewegten und über lange Zeitabschnitte in einem Quadranten standen. Obwohl die Gruppenhaltung für die Mehrzahl der Pferde eine geringere Stressbelastung bedeutete und auch hinsichtlich des Bewegungs- und Sozialverhaltens am artgerechtesten einzustufen ist, sollte für jedes Pferd individuell unter Berücksichtigung seines Alters, seiner Rasse, seines Nutzungsgrades, aber vor allem seines Charakters und seiner Erfahrungen im Sozialverhalten ein passendes großzügig bemessenes Haltungssystem ermittelt werden, um 142 die durch das Haltungssystem ausgeübte Stressbelastung zu minimieren und das Wohlbefinden der Pferde zu steigern. |
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Jankunis, E. S., & Whishaw, I. Q. (2013). Sucrose Bobs and Quinine Gapes: Horse (Equus caballus) responses to taste support phylogenetic similarity in taste reactivity. Behavioural Brain Research, 256, 284–290.
Abstract: Evidence suggests that behavioural affective reactions to sweet and bitter substances are homologous in humans, nonhuman primates, and rodents. The sweet taste of sucrose elicits facial responses that include rhythmic tongue protrusions whereas the bitter taste of quinine elicits facial responses that include gapes, featuring an opening of the mouth and protrusion of the tongue. The present study using the horse (Equus caballus) was undertaken for three reasons: (1) there is debate about the presence of a sweet receptor gene in the horse, (2) there is a need to expand the examination of facial reactions to taste in lineages other than the closely related lineages of rodents and primates, and (3) the horse provides an opportunity to test the hypothesis that some social signals derive from movements related to taste reaction. The horses were given oral infusions of either sucrose or quinine and their behaviour was examined using frame-by-frame video analysis. Control groups were exposed received water or syringe insertion only. Amongst the many responses made to the infusions, the distinctive response to sucrose was a bob coupled with a slight tongue protrusion and forward movement of the ears; the distinctive response to quinine was a head extension and mouth gape accompanied by a large tongue protrusion and backward movement of the ears. Sucrose Bobs and Quinine Gapes are discussed with respect to: (1) the relevance of facial reactions to both sucrose and quinine to taste receptors in horses, (2) the similarity of features of taste expression in horses to those documented in rodents and primates, and (3) the dissimilarity between facial reactions to taste and other social signals displayed by horses.
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Dwan, K., Altman, D. G., Arnaiz, J. A., Bloom, J., Chan, A. - W., Cronin, E., et al. (2008). Systematic Review of the Empirical Evidence of Study Publication Bias and Outcome Reporting Bias. Plos One, 3(8), e3081.
Abstract: Background The increased use of meta-analysis in systematic reviews of healthcare interventions has highlighted several types of bias that can arise during the completion of a randomised controlled trial. Study publication bias has been recognised as a potential threat to the validity of meta-analysis and can make the readily available evidence unreliable for decision making. Until recently, outcome reporting bias has received less attention. Methodology/Principal Findings We review and summarise the evidence from a series of cohort studies that have assessed study publication bias and outcome reporting bias in randomised controlled trials. Sixteen studies were eligible of which only two followed the cohort all the way through from protocol approval to information regarding publication of outcomes. Eleven of the studies investigated study publication bias and five investigated outcome reporting bias. Three studies have found that statistically significant outcomes had a higher odds of being fully reported compared to non-significant outcomes (range of odds ratios: 2.2 to 4.7). In comparing trial publications to protocols, we found that 40-62% of studies had at least one primary outcome that was changed, introduced, or omitted. We decided not to undertake meta-analysis due to the differences between studies. Conclusions Recent work provides direct empirical evidence for the existence of study publication bias and outcome reporting bias. There is strong evidence of an association between significant results and publication; studies that report positive or significant results are more likely to be published and outcomes that are statistically significant have higher odds of being fully reported. Publications have been found to be inconsistent with their protocols. Researchers need to be aware of the problems of both types of bias and efforts should be concentrated on improving the reporting of trials.
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Lagos, L., & Blanco, P. (2021). Testing the use of dogs to prevent wolf attackson free ranging ponies in Iberia? CDPnews, 23, 20–27.
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McLean, A. N., & Christensen, J. W. (2017). The application of learning theory in horse training. Appl. Anim. Behav. Sci., 190, 18–27.
Abstract: The millennia-old practices of horse training markedly predate and thus were isolated from the mid-twentieth century revelation of animal learning processes. From this standpoint, the progress made in the application and understanding of learning theory in horse training is reviewed including a discussion of how learning processes are employed or otherwise under-utilised in training. This review describes the process of habituation and the most commonly applied desensitisation techniques (systematic desensitisation, counter-conditioning, overshadowing, response prevention) and propose two additional techniques (approach conditioning and stimulus blending). The salience of different types of cues, the interaction of operant and classical conditioning and the impact of stress are also discussed. This paper also exposes the inflexibility and occasional inadequacy of the terminology of learning theory when translated from the research laboratory situation to the practical setting in horse training. While learning theory provides a rich toolbox for riders and trainers, the training process is subject to the simultaneous use of multiple learning processes. In addition, learning/behavioural outcomes and trained responses are not just the result of simple stimulus-response based interactions but are further shaped by arousal, affective and attachment states. More research is needed in these areas. For the field of equitation science to progress and to improve clarity and use of learning processes, changes in nomenclature are required. In particular, the use of the terms 'positive' and 'negative' as descriptive labels in both reinforcement and punishment modalities are unacceptably misleading for everyday use. These labels inhibit the understanding and recognition of the learning processes that these terms supposedly represent, yet the learning processes they describe are vital for horse riders, handlers and trainers to understand. We therefore propose that these labels should be re-labelled more appropriately as 'addition' or 'subtraction' reinforcement/punishment. This would enlighten trainers on the correct application of learning theory, and safety and welfare benefits for people and horses would follow. Finally it is also proposed that the term 'conflict theory' be taken up in equitation science to facilitate diagnosis of training-related behaviour disorders and thus enable the emergence of improved training practices. The optimal use of learning theory should be established as a fundamental principle in equestrian education.
Keywords: Behaviour; Conflict theory; Desensitisation; Habituation; Reinforcement; Stress
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Wolter, R. (2018). The behaviour and managementof Przewalski’shorsesin semi-reserves. Ph.D. thesis, University Regensburg, Regensburg.
Abstract: In recent years, Przewalski’s horses have been increasingly kept in semi-reserves. However,there areonly few studies ontheir behaviour and their ability to adaptto management interventions.In the main part of my dissertation, I focus on investigatingthe animals’ behaviour in different semi-reserves with varyinghabitats and living spaces. In addition, I investigate the horses’ behaviour during various management interventionsand analysetheensuing changes instress levels. Another aspect of my dissertation is the studyof social behaviour inPrzewalski’s horses. I investigate theparameters that should be used to demonstrate social bonds between individualsandassess whichdata provide the most meaningful results.In the commentary tochapter 1,several studies investigatingsocial bonds in horsesare discussed. Comparing the various studies, it is strikingthat no homogeneous analyses orevaluation criteria exist. While some authors only considersocial grooming, others include data onthe spatial proximity of the individuals in their evaluations, and various definitionsof proximity can also be found in the literature. Additionally, someauthors use friendly approaches between individuals asa furtherparameter wheninvestigating the social bonds.Continuing with this theme, in chapter 2I investigate the social behaviour of the horses and compare various analysis methods. I show that proactive behaviour, such as friendly approaches, is a good alternative to spatial proximity when investigating social bonds between group members, andis also useful for expanding the often very small data sets of mutual grooming in horses. Comparing Przewalski’s horses with wild living horses, I found no significant differences in the social behavior and the frequency of social interactions, regardlessof group size, group composition, habitat, and individual parameters such as age and gender.Inchapter 3,I investigate the behaviour of a Przewalski’s horse group when exploring a new area of their enclosure. Their behaviour changed, showing less resting and more feeding. Furthermore, the animals maintained greater distances from each other, and the alpha male, instead of herding the group from behind, led the group around the new area and walked in front of the other group members. Moreover, he showed a substantial increase in stress level during the first day.A general comparison of the behaviour of the Przewalski’s horses in different semi-reserves is provided in chapter 4. In it, the habitat choice of the animals and their reactions to various management interventions are investigated. It is shown that Przewalski’s horses prefer open grassland to dense woods, although keeping Przewalski’s horses in a pine forest does not influence the animals’ stress level. In contrast to habitat, food range, and changes in the group composition, which do not appear to change stress levels, individual factors, such as the hierarchy, influence the glucocorticoid level of the animals significantly. The largest increases in stress hormones were demonstrated when the horses were temporarily confined in smaller areas.The importance of the available space is also discussed in chapter 5, where it is shown that horses show less aggressive behaviour when more space is provided. In contrast, the husbandry system does not influence the animals’ aggression, but the way of feeding can additionally reduce agonistic behaviour, for example if food is offered ad libitum.In summary, the results of this study provide indications for the optimization of keeping Przewalski’s horses in semi-reserves. The animals can adapt themselves to the environment and thrive in habitats which do not correspond to their original steppe-like home. Nevertheless, the semi-reserves should provide sufficient grassland, as the horses prefer this type of habitat. General speaking, any types of habitat can only offer a suitable living space if the food range is sufficient for the number of horses. Otherwise, and especially during could winter months, supplementary feeding is necessary according to the body condition of the animals. This is particularly important for older, weakened, or very young animals, which are still adapting to life in the semi-reserve. Without sufficient food, stress hormones can increase and negatively influence the well-being of the horses. The same is true for management interventions: restricting the animals to small enclosures, for example, can adversely affect the horses’ well-being and should be only done if absolutely necessary. Targetedbehaviour observations allow the animals that have a special meaning for the group to be identified, and these should not be taken out of the group unless it is unavoidable, as young and unexperienced horses orientate themselves on those animals. This is especially true for the alpha male in a bachelor group, as these groups are often composed of young horses and the alpha-male provides the necessary stability and experience. Social bonds between individuals can be investigated by observing friendly and proactive behaviour, and social grooming and friendly approaches yield suitable data for such analysis.
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Cozzi, B., Povinelli, M., Ballarin, C., & Granato, A. (2014). The Brain of the Horse: Weight and Cephalization Quotients. Brain Behav Evol, 83(1), 9–16.
Abstract: The horse is a common domestic animal whose anatomy has been studied since the XVI century. However, a modern neuroanatomy of this species does not exist and most of the data utilized in textbooks and reviews derive from single specimens or relatively old literature. Here, we report information on the brain of Equus caballus obtained by sampling 131 horses, including brain weight (as a whole and subdivided into its constituents), encephalization quotient (EQ), and cerebellar quotient (CQ), and comparisons with what is known about other relevant species. The mean weight of the fresh brains in our experimental series was 598.63 g (SEM ± 7.65), with a mean body weight of 514.12 kg (SEM ± 15.42). The EQ was 0.78 and the CQ was 0.841. The data we obtained indicate that the horse possesses a large, convoluted brain, with a weight similar to that of other hoofed species of like mass. However, the shape of the brain, the noteworthy folding of the neocortex, and the peculiar longitudinal distribution of the gyri suggest an evolutionary specificity at least partially separate from that of the Cetartiodactyla (even-toed mammals and cetaceans) with whom Perissodactyla (odd-toed mammals) are often grouped.
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