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Author |
Massen, J.; Sterck, E.; de Vos, H. |
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Title |
Close social associations in animals and humans: functions and mechanisms of friendship |
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Year  |
2010 |
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Behaviour |
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147 |
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11 |
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1379 |
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Both humans and group-living animals associate and behave affiliatively more with some individuals than others. Human friendship has long been acknowledged, and recently scientists studying animal behaviour have started using the term friendship for close social associates in animals. Yet, while biologists describe friends as social tools to enhance fitness, social scientists describe human friendship as unconditional. We investigate whether these different descriptions reflect true differences in human friendship and animal close social associations or are a by-product of different research approaches: namely social scientists focussing on proximate and biologists on ultimate explanations. We first stress the importance of similar measures to determine close social associations, thereafter examine their ultimate benefits and proximate motivations, and discuss the latest findings on the central-neural regulation of social bonds. We conclude that both human friendship and animal close social associations are ultimately beneficial. On the proximate level, motivations for friendship in humans and for close social associations in animals are not necessarily based on benefits and are often unconditional. Moreover, humans share with many animals a similar physiological basis of sociality. Therefore, biologists and social scientist describe the same phenomenon, and the use of the term friendship for animals seems justified. |
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Both humans and group-living animals associate and behave affiliatively more with some individuals than others. Human friendship has long been acknowledged, and recently scientists studying animal behaviour have started using the term friendship for close social associates in animals. Yet, while biologists describe friends as social tools to enhance fitness, social scientists describe human friendship as unconditional. We investigate whether these different descriptions reflect true differences in human friendship and animal close social associations or are a by-product of different research approaches: namely social scientists focussing on proximate and biologists on ultimate explanations. We first stress the importance of similar measures to determine close social associations, thereafter examine their ultimate benefits and proximate motivations, and discuss the latest findings on the central-neural regulation of social bonds. We conclude that both human friendship and animal close social associations are ultimately beneficial. On the proximate level, motivations for friendship in humans and for close social associations in animals are not necessarily based on benefits and are often unconditional. Moreover, humans share with many animals a similar physiological basis of sociality. Therefore, biologists and social scientist describe the same phenomenon, and the use of the term friendship for animals seems justified. |
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Equine Behaviour @ team @ |
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5813 |
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Reyna-Garfias, H.; Miliar, A.; Jarillo-Luna, A.; Rivera-Aguilar, V.; Pacheco-Yepez, J.; Baeza, I.; Campos-Rodríguez, R. |
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Title |
Repeated restraint stress increases IgA concentration in rat small intestine |
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Journal Article |
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Year |
2010 |
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Brain, Behavior, and Immunity |
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24 |
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1 |
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110-118 |
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Restraint-stress; IgA; Small intestine; Polymeric Ig receptor; Catecholamines; Glucocorticoids |
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The most abundant intestinal immunoglobulin and first line of specific immunological defense against environmental antigens is secretory immunoglobulin A. To better understand the effect of repeated stress on the secretion of intestinal IgA, the effects of restraint stress on IgA concentration and mRNA expression of the gene for the alpha-chain of IgA was assessed in both the duodenum and ileum of the rats. Restraint stress induced an increase in intestinal IgA, which was blocked by an adrenalectomy, suggesting a role of catecholamines and glucocorticoids. Whereas the blocking of glucocorticoid receptors by RU-486 did not affect the increased IgA concentration, it did reduce IgA alpha-chain mRNA expression in both segments, indicating a possible mediation on the part of glucocorticoids in IgA secretion by individual cells. Treatment with corticosterone significantly increased both the IgA concentration and IgA alpha-chain mRNA expression in ileum but not in duodenum, suggesting that glucocorticoids may act directly on IgA-antibody forming cells to increase IgA secretion in the former segment. A probable role by catecholamines was evidenced by the reduction in IgA concentration and IgA alpha-chain mRNA expression in both segments after a chemical sympathectomy with 6-hydroxydopamine (6-OHDA). Additionally, norepinephrine significantly reduced IgA alpha-chain mRNA levels but increased pIgR mRNA expression and IgA concentration in both intestinal segments. We propose that the increased intestinal IgA levels caused by repeated restraint stress is likely due to the effects of catecholamines on the transport of plgA across the epithelium. |
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0889-1591 |
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Equine Behaviour @ team @ |
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6002 |
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Peeters, M.; Sulon, J.; Beckers, J.-F.; Ledoux, D.; Vandenheede, M. |
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Comparison between blood serum and salivary cortisol concentrations in horses using an adrenocorticotropic hormone challenge |
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Journal Article |
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Year |
2011 |
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Equine Veterinary Journal |
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43 |
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4 |
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487-493 |
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horse; cortisol; ACTH challenge; saliva; stress |
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Reasons for performing study: In horses, serum cortisol concentration is considered to provide an indirect measurement of stress. However, it includes both free and bound fractions. The sampling method is also invasive and often stressful. This is not the case for salivary cortisol, which is collected using a more welfare-friendly method and represents a part of the free cortisol fraction, which is the biologically active form. Objectives: To compare salivary and serum cortisol assays in horses, in a wide range of concentrations, using an adrenocorticotropic hormone (ACTH) stimulation test, in order to validate salivary cortisol for stress assessment in horse. Methods: In 5 horses, blood samples were drawn using an i.v. catheter. Saliva samples were taken using swabs. Cortisol was assayed by radioimmunoassay. All data were treated with a regression method, which pools and analyses data from multiple subjects for linear analysis. Results: Mean ± s.d. cortisol concentrations measured at rest were 188.81 ± 51.46 nmol/l in serum and 1.19 ± 0.54 nmol/l in saliva. They started increasing immediately after ACTH injection and peaks were reached after 96 ± 16.7 min in serum (356.98 ± 55.29 nmol/l) and after 124 ± 8.9 min in saliva (21.79 ± 7.74 nmol/l, P<0.05). Discharge percentages were also different (225% in serum and 2150% in saliva, P<0.05). Correlation between serum and salivary cortisol concentrations showed an adjusted r2= 0.80 (P<0.001). The strong link between serum and salivary cortisol concentrations was also estimated by a regression analysis. Conclusions: The reliability of both RIAs and regression found between serum and salivary cortisol concentrations permits the validation of saliva-sampling as a noninvasive technique for cortisol level assessment in horses. |
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Blackwell Publishing Ltd |
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2042-3306 |
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Equine Behaviour @ team @ |
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5428 |
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Romero L. M. |
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Using the reactive scope model to understand why stress physiology predicts survival during starvation in Galápagos marine iguanas |
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2011 |
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General and Comparative Endocrinology |
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Gen Comp Endocrinol |
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Reactive scope; Allostasis; Glucocorticoids; Stress; Survival |
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Even though the term “stress” is widely used, a precise definition is notoriously difficult. Notwithstanding this difficulty, stress continues to be an important concept in biology because it attempts to describe how animals cope with environmental change under emergency conditions. Without a precise definition, however, it becomes nearly impossible to make testable a priori predictions about how physiological and hormonal systems will respond to emergency conditions and what the ultimate impact on the animal will be. The reactive scope model is a recent attempt to formulate testable predictions. This model provides a physiological basis to explain why corticosterone negative feedback, but not baseline corticosterone concentrations, corticosterone responses to acute stress, or the interrenal capacity to secrete corticosterone, is correlated with survival during famine conditions in Galápagos marine iguanas. Reactive scope thus provides a foundation for interpreting and predicting physiological stress responses. |
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0016-6480 |
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Equine Behaviour @ team @ |
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5584 |
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Author |
Flauger, B. |
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Title |
The introduction of horses into new social groups with special regard to their stress level |
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2011 |
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Pferd; Equiden; Eingliederungstechnik; Integrationspferd; Stress; Cortisol; Endokrine Reaktion; Gruppenhaltung; Verletzungsgefahr; Aggression; Futterplatzwahl; Kot; Geruchssinn; Mensch-Pferd Interaktion; horse; equids; introduction technique; integration horse; stress; cortisol; endocrine response; group housing; injury risk; aggression; feeding decision; faecal sample; olfaction; human-horse interaction |
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Horses are a highly social species living in complex social systems which should require them to memorise and generalise social experiences and distinguish between familiar and unfamiliar conspecifics. In the main part of my thesis I concentrated on the specific conflict situation of a horse being introduced into a new social group, and investigated its behaviour and stress level. Horses were either introduced (1) immediately, (2) after an observation period, or (3) together with an integration horse after an observation period. Additionally, in the second part of my thesis I arranged several experiments to elaborate additional aspects which could affect the behaviour of horses during introductions. In this study I could describe a simplified method for measuring stress through the analysis of faecal GCMs in horses. An enzyme immunoassay (EIA) for 11-oxoaetiocholanolone using 11-oxoaetiocholanolone-17-CMO: BSA (3?,11-oxo-A EIA) as antigen showed high amounts of immunoreactive substances. The new assay increases the accuracy of the test and lowers the expenses per sample; also storing of samples at room temperature after collection is less critical. This is a big advantage both in the field of wildlife management of equids and in the field of equestrian sports (chapter 1). Comparing the different introduction techniques, the introduction with an integration horse led to significantly less total interactions and lower levels of aggression than the introduction of single horses, both immediately and after several days of observing the new group. Additionally, by observing the behaviour of the horses during everyday sociality I could develop a formula describing the interrelationship between expected aggression level and enclosure size per horse. The curve takes an exponential shape. Starting from a space allowance of 300 m2 and more per horse, the amount of aggressions per hour approaches zero. For the reduction of aggression levels and injury risks in socially kept horses I recommend an enclosure size of at least 300 m2 per horse (chapter 2). I further investigated the stress level of the introduced animals. Horses which were immediately introduced did not show elevated faecal GCMs. In contrast, horses which were introduced after an observation period had slightly elevated values 2 and 3 days after the introduction. For horses introduced together with an integration horse faecal GCMs were significantly above the baseline value on the day of introduction and 1 day after it. These differences between introduction techniques indicate that the introduction event itself is not as stressful as previously assumed. Rather standing together with an integration horse and not being able to integrate immediately into the complete group elicits stress in horses (chapter 3). In the commentary of chapter 4 several studies are discussed which failed to demonstrate social learning in horses. It is argued that they did not consider important aspects which could have an influence, such as the dominance status or the social background of the horses (chapter 4). In chapter 5 a social feeding situation was investigated. The social rank as well as the position of conspecifics affected the feeding strategy of horses. Domestic horses used social cognition and strategic decision making in order to decide where to feed. When possible they tended to return to the same, continuously supplied feeding site and switched to an ?avoidance tendency? in the presence of dominant horses or when another horse was already feeding there (chapter 5). One possibility to recognize group members is through olfactory recognition. In chapter 6 it is shown that horses are able to distinguish their own from their conspecifics? faeces. In addition, they paid most attention to the faeces of those group members from which they received the highest amount of aggressive behaviour (chapter 6). Horses show cognitive abilities because they are able to use humans as local enhancement cues when searching for food, independently of their body posture or gaze consistency when the persons face them. Moreover, they seem to orientate on the attention of familiar persons more than of unfamiliar persons (chapter 7). Altogether, the results of this thesis provide further support for the view that horses show good conflict resolution strategies. They are perfectly able to deal with the conflict situation of being introduced to new group members, and the introduction event itself is not as stressful as previously assumed. It is rather suggested that standing together with an integration horse and not being able to integrate immediately into the complete group elicits stress in horses. All additional experimental set-ups could demonstrate that horses are well capable of social cognition. |
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Ph.D. thesis |
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Equine Behaviour @ team @ epub18463 |
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5736 |
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Author |
Christensen, J.W. |
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Title |
Object habituation in horses: Voluntary vs. negatively reinforced approach to frightening stimuli |
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Conference Article |
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Year |
2012 |
Publication |
Proceedings of the 2. International Equine Science Meeting |
Abbreviated Journal |
Proc. 2. Int. Equine. Sci. Mtg |
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in press |
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Habituation; learning; fearfulness; stress; reinforcement |
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The ability and ease of horses to habituate to frightening stimuli greatly increases safety in the horse-human relationship. Several different techniques have been suggested for habituation training of horses and under certain conditions, preventing animals from avoidance reactions during exposure to frightening stimuli is believed to facilitate habituation. Response prevention does, however, lead to a loss of control, which is a known stress inducer in both animals and humans. This experiment investigated whether horses show increased stress responses when negatively reinforced to approach a mildly frightening stimulus, compared to horses allowed to voluntarily explore the same stimulus. We further investigated whether the prevention of avoidance responses in horses that are negatively reinforced to approach the stimulus, facilitates habituation to the stimulus. Twenty-two 2-3 years old Danish warmblood geldings were included in the study. Half of the horses (NR group) were negatively reinforced (through halter and rope pressure) by a familiar human handler to approach a collection of frightening objects (six open and colourful umbrellas) placed in a semi-circle in a familiar test arena. The other half of the horses were released in the arena and were free to avoid or explore the objects (VOL group). On the next day, all horses were exposed to the objects again without a human to investigate the rate of habituation. Behavioural and heart rate responses were recorded on both days. Data were analysed in a two way repeated measures ANOVA and post hoc analysed via the Holm-Sidak method. In the VOL group, all horses initially chose to avoid the unknown objects, whereas the handler managed to get all horses in the NR group to approach and stand next to the objects within the first 2-min session. As expected, horses in the NR group had a significantly longer duration of alertness (sec, mean ± se: NR: 23 ± 4.1 vs. VOL: 16 ± 4.7, P=0.026) and a higher max HR in the first session (bpm, mean ± se: NR: 106 ± 5.2 vs. VOL: 88 ± 4.4, P=0.004). On the next day, however, the NR horses spent significantly less time investigating the objects (sec, mean ± se: NR: 13 ± 4.1 vs. VOL: 24 ± 6.0, P=0.005) and had a shorter latency to approach a feed container, placed next to the objects (sec, mean ± se: NR: 25 ± 3.9 vs. VOL: 47 ± 16.2, P=0.031), indicating increased habituation. In conclusion, negatively reinforced approach to mildly frightening objects appears to increase stress responses during the initial exposure, but also to facilitate habituation in young horses. |
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Corporate Author |
Christensen, J.W. |
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Xenophon Publishing |
Place of Publication |
Wald |
Editor |
Krueger, K. |
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978-3-9808134-26 |
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Equine Behaviour @ team @ |
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5499 |
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Author |
Flauger, B; Möstl, E.; Krueger., K. |
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Title |
The introduction of horses into new groups: Social interactions and cortisol release |
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Conference Article |
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Year |
2012 |
Publication |
Proceedings of the 2. International Equine Science Meeting |
Abbreviated Journal |
Proc. 2. Int. Equine. Sci. Mtg |
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in press |
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Introduction technique; Aggression; Injury risk; Endocrine response; Stress; Integration horse |
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Domestic horses are kept in so-called “fate societies” where they have to deal with frequent mixing. Several studies have evaluated and discussed the aggression level and injury risk during the introduction of horses into new groups, but nothing is known about the endocrine responses and thus if horses experience stress during introduction.
In this study we analysed the efficiency of four approved introduction techniques and evaluated the introduction of 30 horses into 11 different groups. Horses were introduced: 1) immediately, 2) after observing the new group for several days, 3) together with an “integration horse” after several days of observation, or 4) with a mixed strategy. Aggressive as well as positive social behaviour between the introduced horses and the group members were analysed the two hours following the introduction event. In addition, we focussed on the glucocorticoid production of the newcomer horses by measuring faecal cortisol metabolites (FCM) on the day of the introduction as well as the following three days.
For the four introduction techniques we found significant differences in the horses’ aggressive and submissive behaviour as well as in their total interactions. The introduction together with an integration horse led to significantly lower levels of aggression and less total interactions than the immediate introduction of single horses.
Horses which were introduced immediately or after an observation period showed significantly elevated levels of FCM on the first, second and third day after the introduction. For horses introduced together with an integration horse FCM were already significantly higher on the day of the introduction, indicating a stressful event before the introduction itself. In contrast, FCM levels were always very low when using the mixed technique.
In sum, horses have the ability to deal with conflict when they are introduced to new group members. The introduction event itself appears not to be as stressful as previously assumed. Standing together with an “integration horse” on a separate paddock and not being able to integrate immediately into a new group appears to be stressful for the newcomer. Based on the findings of our study we suggest to introduce new horses in group management together with a new group mate, a so-called “integration horse”. This would reduce the number of total social interactions as well as the aggression level. While this technique may be stressful for the newcomer, it lowers aggressive behaviour between the introduced horse and the group members and consequently reduces injury risks. |
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Corporate Author |
Flauger, B. |
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Xenophon Publishing |
Place of Publication |
Wald |
Editor |
Krueger, K. |
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978-3-9808134-26 |
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Equine Behaviour @ team @ |
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5551 |
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Author |
Hausberger, M.; Fureix, C.; Bourjade, M.; Wessel-Robert, S.; Richard-Yris, M.-A. |
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Title |
On the significance of adult play: what does social play tell us about adult horse welfare? |
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Journal Article |
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Year |
2012 |
Publication |
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Abbreviated Journal |
Naturwissenschaften |
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99 |
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4 |
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291-302 |
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Social play; Stress indicators; Animal welfare; Domestic horse |
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Play remains a mystery and adult play even more so. More typical of young stages in healthy individuals, it occurs rarely at adult stages but then more often in captive/domestic animals, which can imply spatial, social and/or feeding deprivations or restrictions that are challenging to welfare, than in animals living in natural conditions. Here, we tested the hypothesis that adult play may reflect altered welfare states and chronic stress in horses, in which, as in several species, play rarely occurs at adult stages in natural conditions. We observed the behaviour (in particular, social play) of riding school horses during occasional outings in a paddock and measured several stress indicators when these horses were in their individual home boxes. Our results revealed that (1) the number of horses and rates of adult play appeared very high compared to field report data and (2) most stress indicators measured differed between ‘players’ and ‘non-players’, revealing that most ‘playful’ animals were suffering from more chronic stress than ‘non-playful’ horses. Frequency of play behaviour correlated with a score of chronic stress. This first discovery of a relationship between adult play and altered welfare opens new lines of research that certainly deserves comparative studies in a variety of species. |
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Springer-Verlag |
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English |
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0028-1042 |
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Equine Behaviour @ team @ |
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5649 |
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Author |
Christensen, J.W.; Ahrendt, L.P.; Lintrup, R.; Gaillard, C.; Palme, R.; Malmkvist, J. |
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Title |
Does learning performance in horses relate to fearfulness, baseline stress hormone, and social rank? |
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Abstract |
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Year |
2012 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
App Anim Behav Sci |
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Volume |
140 |
Issue |
1 |
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44-52 |
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Horse; Learning; Fearfulness; Stress; Reinforcement; Social rank |
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The ability of horses to learn and remember new tasks is fundamentally important for their use by humans. Fearfulness may, however, interfere with learning, because stimuli in the environment can overshadow signals from the rider or handler. In addition, prolonged high levels of stress hormones can affect neurons within the hippocampus; a brain region central to learning and memory. In a series of experiments, we aimed to investigate the link between performance in two learning tests, the baseline level of stress hormones, measured as faecal cortisol metabolites (FCM), fearfulness, and social rank. Twenty-five geldings (2 or 3 years old) pastured in one group were included in the study. The learning tests were performed by professional trainers and included a number of predefined stages during which the horses were gradually trained to perform exercises, using either negative (NR) or positive reinforcement (PR). Each of the learning tests lasted 3 days; 7min/horse/day. The NR test was repeated in a novel environment. Performance, measured as final stage in the training programme, and heart rate (HR) were recorded. Faeces were collected on four separate days where the horses had been undisturbed at pasture for 48h. Social rank was determined through observations of social interactions during feeding. The fear test was a novel object test during which behaviour and HR were recorded. Performance in the NR and PR learning tests did not correlate. In the NR test, there was a significant, negative correlation between performance and HR in the novel environment (rS=-0.66, P<0.001, i.e. nervous horses had reduced performance), whereas there was no such correlation in the home environment (both NR and PR). Behavioural reactions in the fear test correlated significantly with performance in the NR test in the novel environment (e.g. object alertness and final stage: rS=-0.43, P=0.04), suggesting that performance under unfamiliar, stressful conditions may be predicted by behavioural responses in a fear test. There was a negative correlation between social rank and baseline stress hormones (rS=-0.43, P=0.04), i.e. high rank corresponded to low FCM concentrations, whereas neither rank nor FCM correlated with fearfulness or learning performance. We conclude that performance under stressful conditions is affected by activation of the sympathetic nervous system during training and related to behavioural responses in a standardised fear test. Learning performance in the home environment, however, appears unrelated to fearfulness, social rank and baseline FCM levels. |
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Equine Behaviour @ team @ S0168-1591(12)00168-2 |
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Palme, R. |
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Monitoring stress hormone metabolites as a useful, non-invasive tool for welfare assessment in farm animals |
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2012 |
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Animal Welfare |
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331-337 |
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animal welfare, corticosterone, cortisol, faeces, farm animals, stress |
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A multitude of endocrine mechanisms are involved in coping with challenges. Glucocorticoids, secreted by the adrenal glands, are in the front line of the battle to overcome stressful situations. They are usually measured in plasma samples as parameters of adrenal activity and thus of disturbance. Unfortunately, collecting blood samples itself can disturb an animal. Thus, non-invasive methods for the determination of glucocorticoids or their metabolites have become increasingly popular. The pros and cons of various non-invasive sample materials (saliva, excreta, milk, hair/feathers and eggs) for glucocorticoid determination are given. Above all, faecal samples offer the
advantage that they can be collected easily. In faecal samples, circulating hormone levels are integrated over a certain period of time and represent the cumulative secretion of hormones. Thus, the levels are less affected by short fluctuations or the pulse-like nature of hormone secretion. However, using this technique to assess an animal’s adrenocortical activity is not especially simple. Whether frequent sampling is necessary or single samples will suffice depends upon the study’s aim (whether one is examining the impact of acute or chronic stressors). Background knowledge of the metabolism and excretion of cortisol/corticosterone metabolites is required and a careful validation for each species and sex investigated is obligatory. The present review also addresses analytical issues regarding sample storage, extraction procedures and immunoassays and includes a comprehensive list of published studies (up to 2011) describing the use of such methods in farmed animals. Applied properly, non-invasive techniques to monitor glucocorticoid metabolites in faecal samples of various species are a useful tool for welfare assessment, especially as they are easily applied at farm or group level. |
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