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Beery, A. K., & Kaufer, D. (2015). Stress, social behavior, and resilience: Insights from rodents. Neurobiol. Stress, 1(Stress Resilience), 116–127.
Abstract: The neurobiology of stress and the neurobiology of social behavior are deeply intertwined. The social environment interacts with stress on almost every front: social interactions can be potent stressors; they can buffer the response to an external stressor; and social behavior often changes in response to stressful life experience. This review explores mechanistic and behavioral links between stress, anxiety, resilience, and social behavior in rodents, with particular attention to different social contexts. We consider variation between several different rodent species and make connections to research on humans and non-human primates.
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Christensen, J. W., Beekmans, M., van Dalum, M., & VanDierendonck, M. (2014). Effects of hyperflexion on acute stress responses in ridden dressage horses. Physiol. Behav., 128, 39–45.
Abstract: The effects of hyperflexion on the welfare of dressage horses have been debated. This study aimed to investigate acute stress responses of dressage horses ridden in three different Head-and-Neck-positions (HNPs). Fifteen dressage horses were ridden by their usual rider in a standardised 10-min dressage programme in either the competition frame (CF), hyperflexion (“Low-Deep-and-Round”; LDR) or a looser frame (LF) in a balanced order on three separate test days. Heart rate (HR), heart rate variability parameters (HRV), behaviour and rein tension were recorded during the test. Salivary cortisol concentrations were measured 60min before and 0, 5, 15 and 30min after the test. Rein tension was significantly lower in LF and did not differ between CF and LDR; however approx. 15% of recordings in CF and LDR were above the sensor detection limit of 5kg. The horses had significantly higher cortisol concentrations directly after LDR compared to LF. In addition, the horses showed more distinctive head movements, including head waving, during LDR. There were no significant treatment effects on HR and HRV. In conclusion, the results indicate that LDR may be more stressful to these horses during riding.
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Palme, R., Touma, C., Arias, N., Dominchin, M.N., & Lepschy, M. (2013). Steroid extraction: Get the best out of faecal samples. Wien Tierärztl Monat – Vet Med Austria, 100, 238–246.
Abstract: Faecal steroid hormone metabolites are becoming increasingly popular as parameters for reproductive functions and stress. The extraction of the steroids from the faecal matrix represents the initial step before quantification can be performed. The steroid metabolites present in the faecal matrix are of varying polarity and composition, so selection of a proper extraction procedure is essential. There have been some studies to address this complex but often neglected point. Radiolabelled steroids (e.g. cortisol or progesterone) have frequently been added to faecal samples to estimate the efficiency of the extraction procedures used. However, native, unmetabolized steroids are normally not present in the faeces and therefore the results are artifi- cial and do not accurately reflect the actual recoveries of the substances of interest. In this respect, recovery experiments based on faecal samples from radiometabolism studies are more informative. In these samples, the metabolite content accurately reflects the mixture of metabolites present in the given species. As a result, it is possible to evaluate different extraction methods for use with faecal samples. We present studies on sheep, horses, pigs, hares and dogs that utilized samples containing naturally metabolized, 14C-labelled steroids. We recommend extracting faecal steroids by simply suspending the faeces in a high percentage of a primary alcohol (for glucocorticoid metabolites 80% aqueous methanol proved best suited for virtually all mammalian species tested so far). Not only does the procedure significantly increase the total amount of recovered radioactivity, it also increases the percentage of unconjugated metabolites, which are more likely to be recognized by the antibodies used in various immunoassays. The advantages of this extraction procedure are clear: it is very easy to use (no evaporation step is needed), it yields high recoveries and variation based on the extraction procedure is reduced to a minimum.
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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.
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Krueger, K., Esch, L., Farmer, K., & Marr, I. (2021). Basic Needs in Horses?--A Literature Review. Animals, 11(6), 1798.
Abstract: Every animal species has particular environmental requirements that are essential for its welfare, and when these so-called “basic needs” are not fulfilled, the animals suffer. The basic needs of horses have been claimed to be social contact, social companionship, free movement and access to roughage. To assess whether horses suffer when one or more of the four proposed basic needs are restricted, we examined several studies (n = 38) that reported behavioural and physiological reactions to these restrictions. We assigned the studies according to the four types of responses investigated: (a) Stress, (b) Active, (c) Passive, and (d) Abnormal Behaviour. Furthermore, the number of studies indicating that horses reacted to the restrictions were compared with the number of studies reporting no reaction. The limited number of studies available on single management restrictions did not allow conclusions to be drawn on the effect of each restriction separately, especially in the case of social companionship. However, when combinations of social contact, free movement and access to roughage were restricted, many of the horses had developed responses consistent with suffering. Passive Responses, indicating acute suffering, and Abnormal Behaviour, indicating suffering currently or at some time in the past, were especially clearly demonstrated. This provides further evidence of the usefulness of assessing behavioural parameters in combination with physiological measurements when evaluating horse welfare. This meta-analysis of the literature confirms that it is justified to claim that social contact, free movement and access to roughage are basic needs in horses.
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Krueger, K., Schwarz, S., Marr, I., & Farmer, K. (2022). Laterality in Horse Training: Psychological and Physical Balance and Coordination and Strength Rather Than Straightness. Animals, 12(8), 1042.
Abstract: For centuries, a goal of training in many equestrian disciplines has been to straighten the horse, which is considered a key element in achieving its responsiveness and suppleness. However, laterality is a naturally occurring phenomenon in horses and encompasses body asymmetry, motor laterality and sensory laterality. Furthermore, forcibly counterbalancing motor laterality has been considered a cause of psychological imbalance in humans. Perhaps asymmetry and laterality should rather be accepted, with a focus on training psychological and physical balance, coordination and equal strength on both sides instead of enforcing “straightness”. To explore this, we conducted a review of the literature on the function and causes of motor and sensory laterality in horses, especially in horses when trained on the ground or under a rider. The literature reveals that body asymmetry is innate but does not prevent the horse from performing at a high level under a rider. Motor laterality is equally distributed in feral horses, while in domestic horses, age, breed, training and carrying a rider may cause left leg preferences. Most horses initially observe novel persons and potentially threatening objects or situations with their left sensory organs. Pronounced preferences for the use of left sensory organs or limbs indicate that the horse is experiencing increased emotionality or stress, and long-term insufficiencies in welfare, housing or training may result in left shifts in motor and sensory laterality and pessimistic mentalities. Therefore, increasing laterality can be regarded as an indicator for insufficiencies in housing, handling and training. We propose that laterality be recognized as a welfare indicator and that straightening the horse should be achieved by conducting training focused on balance, coordination and equal strength on both sides.
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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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Abeyesinghe, S. M., Nicol, C. J., Wathes. C.M., & Randall, J. M. (2001). Development of a raceway method to assess aversion of domestic fowl to concurrent stressors. Behav. Process., 56(3), 175–194.
Abstract: The requirement for assessing the effects of stressor combinations in improving the welfare of animals has not been widely recognised. Knowledge of the effects of concurrent stressors is needed to improve environments such as transport, where animals are presented with many simultaneous challenges. However, no method for measuring the effects of different stressors with a common unit is currently available. A locomotor passive avoidance method was developed as a common currency measure of the aversion of domestic fowl to concurrent stressors, using vibrational and thermal stressors as an exemplar. Juvenile fowl, fasted overnight, were trained to run a raceway into a goal-box for small food rewards (FR1). When running consistently, the reinforcement schedule was superimposed with a FR5 treatment schedule (60 min confinement in the goal-box with either a control of no other stressors [N] or concurrent vibration and thermal stressors [VT]). Subsequent latency to return to the goal-box was recorded as a measure of aversion. The factors affecting bird response were addressed in a series of experiments to optimise the method and clarify interpretation of results. Pre-feeding (20% ration 2 h prior to testing) did not affect response, but increasing the number of treatment presentations facilitated learning and increased method sensitivity. Treatment responses were consistent across experiments; overall VT was avoided (P<0.001), but N was not. However, there was large individual variation in response to VT. A final experiment indicated that, given a visual discriminatory cue, birds were capable of learning the required association between entering the goal-box and receiving the treatment, suggesting that the delay responses were due to aversion rather than the immediate impact of treatment on ability to respond. Further work is required to test the singular stressors, but the method retains common currency potential for assessing aversion to multiple stressors.
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Overli, O., Korzan, W. J., Hoglund, E., Winberg, S., Bollig, H., Watt, M., et al. (2004). Stress coping style predicts aggression and social dominance in rainbow trout. Horm Behav, 45(4), 235–241.
Abstract: Social stress is frequently used as a model for studying the neuroendocrine mechanisms underlying stress-induced behavioral inhibition, depression, and fear conditioning. It has previously been shown that social subordination may result in increased glucocorticoid release and changes in brain signaling systems. However, it is still an open question which neuroendocrine and behavioral differences are causes, and which are consequences of social status. Using juvenile rainbow trout of similar size and with no apparent differences in social history, we demonstrate that the ability to win fights for social dominance can be predicted from the duration of a behavioral response to stress, in this case appetite inhibition after transfer to a new environment. Moreover, stress responsiveness in terms of confinement-induced changes in plasma cortisol was negatively correlated to aggressive behavior. Fish that exhibited lower cortisol responses to a standardized confinement test were markedly more aggressive when being placed in a dominant social position later in the study. These findings support the view that distinct behavioral-physiological stress coping styles are present in teleost fish, and these coping characteristics influence both social rank and levels of aggression.
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Palagi, E., Cordoni, G., & Borgognini Tarli, S. (2006). Possible roles of consolation in captive chimpanzees (Pan troglodytes). Am J Phys Anthropol, 129(1), 105–111.
Abstract: Empathy is a necessary prerequisite for the occurrence of consolation. The term “consolation” contains a hypothesis about function, which is distress alleviation. The present study aims to confirm the occurrence of consolation in captive chimpanzees via the post-conflict/matched-control method (PC-MC) and to suggest its possible roles. We collected 273 PC-MC pairs in the group of Pan troglodytes housed in the ZooParc de Beauval (France). We confirmed the presence of consolatory contacts (mean level of consolation, 49.5% +/- 22.3% SEM) in the colony. Consolation rates were significantly higher than reconciliation levels (mean level of reconciliation, 28.9% +/- 16.8% SEM). The level of consolation was greater in the absence of reconciliation than in the presence of it, suggesting that consolation might be an alternative behavior. As friendship and relatedness did not influence the occurrence of consolation, they did not seem to be the best prerequisites for this behavioral mechanism, at least in this chimpanzee colony. Affinitive contacts with third parties were significantly more frequent when the victim called attention to itself during severe aggressions by screaming. These high-pitched sounds seem to be useful in eliciting aid from conspecifics, as occurs in young humans. The occurrence of consolation reduced the likelihood of further attacks among group-members. From this perspective, both victims and consolers most likely gain potential advantages by interacting with each other when aggression is particularly severe, reconciliation is not immediate, and consequently social stress reaches high levels.
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