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h:, M.; Lévy, F.; Fortin, M.; Leterrier, C.; LansadLansade, L. |
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Stress and temperament affect working memory performance for disappearing food in horses, Equus caballus |
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2013 |
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Animal Behaviour |
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86 |
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6 |
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1233-1240 |
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delayed-response task emotion; equid; Equus caballus; fearfulness; individual difference; personality; stress; temperament |
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In the present study, we sought to determine the influence of stress and temperament on working memory for disappearing food in horses. After assessment of five dimensions of temperament, we tested working memory of horses using a delayed-response task requiring a choice between two food locations. Delays ranging from 0 to 20 s were tested. The duration of working memory for disappearing food was first characterized without stressors (N = 26). The horses were then divided into two groups and their performance was assessed under stressful (exposure to acute stressors prior to testing, N = 12) or control conditions (N = 12). Results showed that the duration of working memory for disappearing food lasted at least 20 s under nonstressful conditions, and that under stressful conditions this duration lasted less than 12 s. This stress-induced impairment confirms in a nonrodent species that working memory performance is very sensitive to exposure to stressors. In addition, working memory performance in horses is influenced by the temperamental dimension of fearfulness according to the state of stress: fearful horses showed better performance under control conditions and worse performance under stressful conditions than nonfearful horses. These findings are discussed in the context of the Yerkes–Dodson law of stress and performance. |
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0003-3472 |
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Equine Behaviour @ team @ |
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5746 |
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Dreschel, N.A.; Granger, D.A. |
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Title |
Methods of collection for salivary cortisol measurement in dogs |
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Journal Article |
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Year |
2009 |
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Hormones and Behavior |
Abbreviated Journal |
Horm. Behav. |
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55 |
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1 |
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163-168 |
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Dog; Canine; Salivary cortisol; Methods; Measurement; Stress |
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Salivary cortisol has been increasingly used as a measure of stress response in studies of welfare, reaction to stress and human–animal interactions in dogs and other species. While it can be a very useful measure, there are a number of saliva collection issues made evident through studies in the human and animal fields which have not been investigated in the canine species. Collection materials and the volume of saliva that is collected; the use of salivary stimulants; and the effect of food contamination can all dramatically impact cortisol measurement, leading to spurious results. In order to further examine the limitations of the collection method and the effects of collection material and salivary stimulant on salivary cortisol levels, a series of clinical, in vitro and in vivo studies were performed. It was found that there is a large amount of inter- and intra-individual variation in salivary cortisol measurement. Beef flavoring of collection materials leads to unpredictable variability in salivary cortisol concentration. Using salivary stimulants such as citric acid also has the potential to affect cortisol concentration measurement in saliva. Hydrocellulose appears to be a useful collection material for salivary cortisol determination. Recommendations for collection materials and use of salivary stimulants are presented. |
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Equine Behaviour @ team @ |
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5560 |
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Romero, L.M.; Dickens, M.J.; Cyr, N.E. |
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Title |
The reactive scope model — A new model integrating homeostasis, allostasis, and stress |
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Journal Article |
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Year |
2009 |
Publication |
Hormones and Behavior |
Abbreviated Journal |
Horm. Behav. |
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55 |
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3 |
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375-389 |
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Keywords |
Stress; Allostasis; Glucocorticoids; Fight-or-flight; Homeostasis |
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Allostasis, the concept of maintaining stability through change, has been proposed as a term and a model to replace the ambiguous term of stress, the concept of adequately or inadequately coping with threatening or unpredictable environmental stimuli. However, both the term allostasis and its underlying model have generated criticism. Here we propose the Reactive Scope Model, an alternate graphical model that builds on the strengths of allostasis and traditional concepts of stress yet addresses many of the criticisms. The basic model proposes divergent effects in four ranges for the concentrations or levels of various physiological mediators involved in responding to stress. (1) Predictive Homeostasis is the range encompassing circadian and seasonal variation — the concentrations/levels needed to respond to predictable environmental changes. (2) Reactive Homeostasis is the range of the mediator needed to respond to unpredictable or threatening environmental changes. Together, Predictive and Reactive Homeostasis comprise the normal reactive scope of the mediator for that individual. Concentrations/levels above the Reactive Homeostasis range is (3) Homeostatic Overload, and concentrations/levels below the Predictive Homeostasis range is (4) Homeostatic Failure. These two ranges represent concentrations/levels with pathological effects and are not compatible with long-term (Homeostatic Overload) or short-term (Homeostatic Failure) health. Wear and tear is the concept that there is a cost to maintaining physiological systems in the Reactive Homeostasis range, so that over time these systems gradually lose their ability to counteract threatening and unpredictable stimuli. Wear and tear can be modeled by a decrease in the threshold between Reactive Homeostasis and Homeostatic Overload, i.e. a decrease in reactive scope. This basic model can then be modified by altering the threshold between Reactive Homeostasis and Homeostatic Overload to help understand how an individual's response to environmental stressors can differ depending upon factors such as prior stressors, dominance status, and early life experience. We illustrate the benefits of the Reactive Scope Model and contrast it with the traditional model and with allostasis in the context of chronic malnutrition, changes in social status, and changes in stress responses due to early life experiences. The Reactive Scope Model, as an extension of allostasis, should be useful to both biomedical researchers studying laboratory animals and humans, as well as ecologists studying stress in free-living animals. |
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0018-506x |
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Equine Behaviour @ team @ |
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5583 |
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Author |
Romero L. M. |
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Title |
Using the reactive scope model to understand why stress physiology predicts survival during starvation in Galápagos marine iguanas |
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Journal Article |
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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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Sloet van Oldruitenborgh-Oosterbaan, M.M.; Blok, M.B.; Begeman, L.; Kamphuis, M.C.D.; Lameris, M.C.; Spierenburg, A.J.; Lashley, M.J.J.O. |
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Workload and stress in horses: comparison in horses ridden deep and round ('rollkur') with a draw rein and horses ridden in a natural frame with only light rein contact |
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Journal Article |
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Year |
2006 |
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Tijdschrift Voor Diergeneeskunde |
Abbreviated Journal |
Tijdschr Diergeneeskd |
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131 |
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5 |
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152-157 |
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Animal Husbandry/methods; Animals; Biomechanics; Blood Glucose/analysis; Female; Heart Rate/physiology; Hematocrit/veterinary; Horses/blood/*physiology; Hydrocortisone/blood; Lactic Acid/blood; Physical Conditioning, Animal/adverse effects/*physiology; Stress, Physiological/blood/etiology/veterinary |
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'Rollkur' or 'overbending' is the low and deep riding of a dressage horse during training or warming up. Lately, this technique has been criticized, and not necessarily objectively, on welfare grounds. To be able to evaluate these criticisms, more needs to be known about the workload and stress of horses being ridden 'rollkur'. The aim of the present study was to compare the workload of eight riding-school horses when being ridden deep and round with a draw rein ('rollkur') and when being ridden in a natural frame with only light rein contact ('free'). Workload (as measured by heart rate and blood lactate concentration) was slightly higher when horses were ridden 'rollkur' than when they were ridden 'free'. There were no differences in packed cell volume, or glucose and cortisol concentrations. No signs of uneasiness or stress could be determined when the horses were ridden 'rollkur'. Subjectively, all horses improved their way of moving during 'rollkur' and were more responsive to their rider. |
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Department of Equine Sciences, Faculty of Veterinary Medicine, Utrecht University, Yalelaan 12, 3584 CM Utrecht, The Netherlands. m.sloet@vet.uu.nl |
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0040-7453 |
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PMID:16532786 |
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Equine Behaviour @ team @ |
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5638 |
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Author |
Innes, L.; McBride, S. |
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Negative versus positive reinforcement: An evaluation of training strategies for rehabilitated horses |
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Journal Article |
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2008 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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112 |
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3–4 |
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357-368 |
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Horse; Training; Positive reinforcement; Negative reinforcement; Stress; Rehabilitation |
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Rescued equids are often exposed to rehabilitation and training (or retraining) programmes to improve their physical and psychological well-being as well as to facilitate the re-homing process. Training uses either positive or negative reinforcement learning procedures and it is considered here that, there may be welfare implications associated with using the latter technique as it has the potential to overlay acute stress on animals with a chronic stress life history. The aim of this study, therefore, was to compare these training strategies (negative versus positive reinforcement) on equine behaviour and physiology as the first step in establishing an optimal rehabilitation approach (from a welfare perspective) for equids that have been subjected to chronic stress in the form of long-term neglect/cruelty. Over a 7-week period, 16 ponies (aged 6–18 months) were trained using either positive (‘positive’) (n = 8) or negative reinforcement (‘negative’) (n = 8) techniques to lead in hand, stand to be groomed, traverse an obstacle course and load into a trailer. Heart rate was measured (5 s intervals) on days 1 and 4 of each training week, ‘Pre’- (1 h), ‘During’ (0.5 h) and ‘Post’- (1 h) training session. Ethograms (10.00–20.00 h) outside of the training period were also compiled twice weekly. In addition, weekly arena tests (as a measure of reactivity) were also performed 1 week before and during the 7 weeks of training. Results showed significant differences between the two training schedules for some measures during the latter stages of the trial and suggested that animals trained under a positive reinforcement schedule were more motivated to participate in the training sessions and exhibited more exploratory or ‘trial and error’ type behaviours in novel situations/environments. In this context, the incorporation of positive reinforcement schedules within a rehabilitation programme may be of benefit to the animal from a welfare perspective. |
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0168-1591 |
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Equine Behaviour @ team @ |
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5644 |
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Author |
Hausberger, M.; Fureix, C.; Bourjade, M.; Wessel-Robert, S.; Richard-Yris, M.-A. |
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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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2012 |
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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 |
Bosch, O.J.; Nair, H.P.; Ahern, T.H.; Neumann, I.D.; Young, L.J. |
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Title |
The CRF System Mediates Increased Passive Stress-Coping Behavior Following the Loss of a Bonded Partner in a Monogamous Rodent |
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Journal Article |
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2008 |
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Neuropsychopharmacology |
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34 |
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6 |
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1406-1415 |
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prairie vole; passive stress-coping; forced swim test; tail suspension test; elevated plus-maze; hypothalamo–pituitary–adrenal axis |
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Social relationships significantly influence physiology and behavior, including the hypothalamo–pituitary–adrenal axis, anxiety, and mental
health. Disruption of social bonds through separation or death often results in profound grieving, depression, and physical illness. As the
monogamous prairie vole forms enduring, selective pair bonds with the mating partner, they provide an animal model to study the
physiological consequences of pair bonding and, thus, the loss of the bonded partner. Male prairie voles were paired with a novel female
or male sibling. After 5 days, half of the males of each group were separated from the partner. Elevated plus-maze, forced swim, and tail
suspension tests were used to assess anxiety-like and passive stress-coping behaviors indicative of depressive-like behavior. Following 4
days of separation from the female but not the male partner, experimental males displayed increased passive stress-coping. This effect
was abolished by long-term intracerebroventricular infusion of a nonselective corticotropin-releasing factor (CRF) receptor antagonist
without disrupting the bond itself. Both CRF type 1 and 2 receptors were involved in the emergence of passive stress-coping behavior.
Furthermore, pairing with a female was associated with elevated CRF mRNA in the bed nucleus of the stria terminalis, and partner loss
elicited a pronounced increase in circulating corticosteroid and adrenal weight. We speculate that the CRF system may mediate an
aversive affect following separation from the female partner, which may facilitate proximity seeking between the pair-bonded individuals.
Hence, the prairie vole model may provide insights into brain mechanisms involved in the psychopathological consequences of partner
loss. |
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American College of Neuropsychopharmacology |
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0893-133x |
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Equine Behaviour @ team @ |
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5731 |
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Author |
Flauger, B. |
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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.; Ahrendt, L.P.; Lintrup, R.; Gaillard, C.; Palme, R.; Malmkvist, J. |
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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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140 |
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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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0168-1591 |
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Equine Behaviour @ team @ S0168-1591(12)00168-2 |
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5769 |
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