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Ostner, J., Heistermann, M., & Schülke, O. (2008). Dominance, aggression and physiological stress in wild male Assamese macaques (Macaca assamensis). Hormones and Behavior, 54(5), 613–619.
Abstract: In group-living animals relative rank positions are often associated with differences in glucocorticoid output. During phases of social stability, when dominance positions are clear and unchallenged, subordinates often face higher costs in terms of social stress than dominant individuals. In this study we test this prediction and examine additional potential correlates of stress, such as reproductive season, age and amount of aggression received in wild, seasonally breeding Assamese macaques (Macaca assamensis). During a mating and a non-mating season we collected 394 h of focal observational data and 440 fecal samples of six adult and six large subadult males living in a multimale-multifemale group in their natural habitat in northeastern Thailand. The mating season was characterized by a general increase in aggressive behavior and glucocorticoid excretion across all males compared to the non-mating season. Among adult males, mating season glucocorticoid levels were significantly negatively related with dominance rank and positively with the amount of aggression received. Both relationships were non-significant among large subadult males. Thus, our results suggest that in adult Assamese macaques a high dominance position is not associated with high costs. Low costs of dominance might be induced by strong social bonds among top-ranking males, which exchange frequent affiliative interactions and serve as allies in coalitionary aggression against potentially rank-challenging subordinate males.
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Nagy, K., Bodó, G., Bárdos, G., & Harnos, A. (2008). Is modified Forssell"s operation superior to cribbing collar in preventing crib-biting in horses? In IESM 2008.
Abstract: Crib-biting (wind-sucking) might be a coping response of the horses to the challenges of
uncontrolled environmental events. Prevention of this stereotypic behaviour evokes physiological responses consistent with increased stress. Reducing the incidence of cribbiting, however, is important in order to prevent undesirable physical and behavioural consequences (tooth erosion, altered gut function, gastric inflammation/ulceration, colic, etc.). Common treatment of crib-biting is the application of a cribbing collar, which limits the flexion of the neck making this stereotypic movement uncomfortable and difficult. Another method, the modified Forssell"s operation, is becoming more and more popular amongst the horse owners. It is based on the removal of the muscles used in crib-biting (m.omohyoideus, m.sternohyoideus, m.sternothyrohyoideus) and the ventral branches of the spinal accessory nerves. Surveys on the success of this surgical procedure have revealed inconsistent results, and, contrary to the cribbing collar, its effect on the stress level have not been studied either. The aim of our study was to determine whether the modified Forssell"s procedure is superior to the cribbing collar treatment. Differences in stress management was tested by a crib-biting provoking test, in which surgically treated horses, crib-biting horses, crib-biting horses with cribbing collar, and normal horses (those showing no stereotypies), altogether 56 horses were compared. In this test, a food bucket had been placed out of the reach of the animal, from which titbits were given 3 times. Behaviour and heart rate variability (HRV) of the horses were recorded and analysed throughout the test. Hypotheses were tested by linear mixed model. According to our results, both prevention methods (collar or surgery) inhibited crib-biting successfully though not totally. Regarding behaviour and heart rate variability, horses prevented from crib-biting (by collar or surgery) differed significantly from crib-biting and normal horses but not from each other. Normal horses were usually trying to reach the food-bucket while present and were standing still afterwards, whereas the other three groups had not really made efforts to reach the bucket, spent less time with resting, and performed or tried crib-biting. During the stress-test, normal and crib-biting horses had shown good stress-adaptation to the challenge since their HRV, after an initial increase, returned to the basal value by the end. On the contrary, HRV of the two prevented groups remained elevated and showed large oscillations throughout. They had not found a successful coping behaviour either. Our results suggest that since prevention may significantly increase distress, the treatment in itself, without changing the motivation of the horse to perform the replacement behaviour – it seems to be unsatisfactory and insufficient. After prevention the motivation of the horse to perform crib-biting should be addressed. In addition, considering that prevention by collar and surgery had not resulted in any significant behavioural or physiological differences, the superiority of the modified Forssell"s procedure might be questioned. However, the surgery might be recommended if treatment with collar is ineffective. |
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Nagy, K., Bodó, G., Bárdos, G., Harnos, A., & Kabai, P. (2009). The effect of a feeding stress-test on the behaviour and heart rate variability of control and crib-biting horses (with or without inhibition). Appl. Anim. Behav. Sci., 121(2), 140–147.
Abstract: Crib-biting is a form of oral stereotypy affecting 4-5% of horses. Once fixed, crib-biting is difficult to eliminate by behaviour therapy, however, its performance can be inhibited by collar or surgery treatment (modified Forssell's procedure). Although surgical intervention is widespread, the effects on stress coping in horses have not been studied. In the present study we evaluated changes in behaviour response and heart rate variability in 9 control, 10 crib-biting, 10 collar and 11 surgically treated horses in a feeding stress-test, in which a feeding-bowl was placed in front but out of the reach of the horses, from which tidbits were given 3 times. We found that stress triggers high oral activity, mainly cribbing in crib-biting horses, elevates other forms of oral activities in the inhibited groups and does not affect oral activities of controls. Instead of performing oral activities, control horses tended to target an unavailable feeding-bowl by pawing or head-tossing. Changes in stress level were indistinguishable in controls and crib-biters as heart rate variability returned to baseline values in both groups. In contrast, horses inhibited to perform crib-biting showed elevated stress level throughout the test period. Our results suggest that crib-biting may develop to cope with stress, and such coping function diminishes when inhibited.
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Carlstead, K., & Brown, J. L. (2005). Relationships between patterns of Fecal corticoid excretion and behavior, reproduction, and environmental factors in captive black (Diceros bicornis) and white (Ceratotherium simum) rhinoceros. Zoo Biol., 24(3), 215–232.
Abstract: Mortality is high in zoo-housed black rhinoceros (Diceros bicornis), and the reproductive rates of captive white rhinoceros (Ceratotherium simum) are unsustainably low. To determine the possible role of stress in the causation of these problems, we analyzed weekly fecal samples collected for 1 year from black (10 males and 16 females) and white (six males and 13 females) rhinoceroses at 16 zoos for corticoid metabolite concentrations. Fecal corticoid profiles were examined in relation to behavior as rated by keepers in a questionnaire, luteal phase ovarian cycles of females (Brown et al., 2001), and socioenvironmental factors. We compared individual fecal corticoid profiles by examining hormone means and variability (i.e., standard deviation (SD) and coefficient of variation (CV)). For the black rhinos, higher mean corticoid concentrations were found at zoos where rhinos were maintained in enclosures that were exposed to the public around a greater portion of the perimeter. Higher variability in corticoid excretion was correlated with higher rates of fighting between breeding partners and higher institutional mortality rates. Black rhino pairs that were kept separated exhibited lower corticoid variability and less fighting activity when they were introduced during female estrous periods compared to pairs that were kept together every day. For white rhinos, significantly lower mean corticoids were found for individuals that rated higher on “friendliness to keeper.” Higher corticoid variability was found in noncycling as compared to cycling white rhino females. Noncycling females exhibited higher rates of stereotypic pacing and lower frequencies of olfactory behaviors. Interindividual differences in mean corticoids in both species appeared to be related to responsiveness to humans, whereas corticoid variability was related to intraspecific social relationships. More importantly, high corticoid variability appeared to be an indicator of chronic or “bad” stress, because of its association with potentially deleterious consequences in each species (i.e., fighting and mortality (black rhino), and reproductive acyclicity (white rhino)). Our results provide evidence that social stressors may cause chronic stress in black and white rhinos, and that this contributes to the captive-population sustainability problems observed in each species. Zoo Biol 0:1–18, 2005. © 2005 Wiley-Liss, Inc.
Keywords: stress; adrenal activity; olfactory behavior; ovarian activity; reproduction; mortality
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Romero, L. M., Dickens, M. J., & Cyr, N. E. (2009). The reactive scope model — A new model integrating homeostasis, allostasis, and stress. Horm. Behav., 55(3), 375–389.
Abstract: 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.
Keywords: Stress; Allostasis; Glucocorticoids; Fight-or-flight; Homeostasis
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Mormède, P., Andanson, S., Aupérin, B., Beerda, B., Guémené, D., Malmkvist, J., et al. (2007). Exploration of the hypothalamic-pituitary-adrenal function as a tool to evaluate animal welfare. Physiol. Behav., 92(3), 317–339.
Abstract: Measuring HPA axis activity is the standard approach to the study of stress and welfare in farm animals. Although the reference technique is the use of blood plasma to measure glucocorticoid hormones (cortisol or corticosterone), several alternative methods such as the measurement of corticosteroids in saliva, urine or faeces have been developed to overcome the stress induced by blood sampling itself. In chronic stress situations, as is frequently the case in studies about farm animal welfare, hormonal secretions are usually unchanged but dynamic testing allows the demonstration of functional changes at several levels of the system, including the sensitization of the adrenal cortex to ACTH and the resistance of the axis to feedback inhibition by corticosteroids (dexamethasone suppression test). Beyond these procedural aspects, the main pitfall in the use of HPA axis activity is in the interpretation of experimental data. The large variability of the system has to be taken into consideration, since corticosteroid hormone secretion is usually pulsatile, follows diurnal and seasonal rhythms, is influenced by feed intake and environmental factors such as temperature and humidity, age and physiological state, just to cite the main sources of variation. The corresponding changes reflect the important role of glucocorticoid hormones in a number of basic physiological processes such as energy metabolism and central nervous system functioning. Furthermore, large differences have been found across species, breeds and individuals, which reflect the contribution of genetic factors and environmental influences, especially during development, in HPA axis functioning. Usually, these results will be integrated with data from behavioral observation, production and pathology records in a comprehensive approach of farm animal welfare.
Keywords: Stress; Animal welfare; HPA axis; Glucocorticoid hormones; Acth; Dexamethasone suppression test; Cattle; Pig; Fur animals; Mink; Fox; Poultry; Fish
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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.
Keywords: Stress; Anxiety; Social behavior; Sociality; Social stress; Social buffering
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Mendl, M. (1999). Performing under pressure: stress and cognitive function. Appl. Anim. Behav. Sci., 65(3), 221–244.
Abstract: The way in which cognitive functioning is affected by stressors is an important area of research for applied ethologists because stress caused by captive conditions may disrupt cognitive processes and lead to welfare and husbandry problems. Such problems may be minimised through an understanding of the links between stress and cognition. The effects of stress on cognitive function have been studied in disciplines ranging from human perceptual psychology to animal neuroscience. The aim of this paper is to provide an introduction to this research, focusing on the effects of stressors on attention, memory formation and memory recall. Findings from such a diverse literature with little apparent inter-disciplinary communication are inevitably complex and often contradictory. Nevertheless, some generalities do emerge. The idea that an inverted U-shaped relationship exists between an individual's state of stress or arousal and its ability to perform a cognitive task effectively, the so-called Yerkes-Dodson law, is commonly encountered. The law has limited explanatory value because it is unlikely that different stressors act on cognitive function via the same intervening, non-specific state. Furthermore, the law only provides a very general description of the relationship between stress and cognitive function. Empirical research on attention and memory processes reveals more specific findings. Stressors appear to cause shifts, lapses and narrowing of attention, and can also influence decision speed. These processes may be viewed as serving an adaptive role helping the animal to search for and scrutinise a source of danger. There is conflicting evidence as to whether hormones involved in the hypothalamic-pituitary-adrenal stress response play a part in these processes. These hormones and those involved in the sympathetic-adrenomedullary stress response do appear to play an important role in memory formation. Low or moderate concentrations of circulating glucocorticoids and catecholamines can enhance memory formation, while excessively high or prolonged elevations of these hormones can lead to memory disruption. The effects of stressors on memory recall are less clear. There is evidence for disruptive effects, and for facilitatory effects indicating state-dependent memory recall; events experienced under conditions of high arousal may be best recalled under similar conditions. Applied ethologists have the opportunity to extend work in this area, which often involves studies of single stressors/stress hormones acting in isolation and limited measures of cognitive function, by focusing on real-life husbandry stressors encountered by captive animals. This will yield fundamental information which also has direct relevance to animal welfare and management issues.
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König v. Borstel, U., Visser, E. K., & Hall, C. (2017). Indicators of stress in equitation. Appl. Anim. Behav. Sci., 190, 43–56.
Abstract: Abstract Stress is a generic concept describing the body's reaction to external stimuli, including both physiological and psychological factors. Therefore, by definition, the assessment of psychological stress in the exercising horse encompasses the problem of teasing apart the psychological and physiological factors both of which result in stress responses. The present study reviews the existing literature on various measures of stress taken specifically in the context of equitation science. Particular attention has been paid to short-term effects, and commonly used measurements of short-term stress include heart rate, a number of heart rate variability parameters, blood or saliva cortisol levels, eye temperature, and various behaviour parameters including in particular behaviour patterns presumably indicative of conflict with the rider's/trainer's aids. Inspection of the individual studies' results revealed that disagreement between these different measures of stress is commonplace. For physiological parameters, the largest proportion of agreement (i.e. both parameters simultaneously indicated either higher, insignificant or lower stress compared to a control treatment) was found for heart rate and heart rate variability parameters, while generally limited agreement was found for cortisol. It appears that cortisol levels may not be particularly useful for assessing/assessment of the valence of a situation in the exercising horse as cortisol levels are predominantly linked to activation and exercise levels. Although heart rate variability parameters reflect in theory more closely sympathovagal balance compared to cortisol levels, great care has to be taken regarding the use of appropriate time-frames, appropriate raw data correction methods as well as the use of appropriate equipment. In spite of its wide-spread and apparently successful use, popular equipment may in fact not be accurate enough under field conditions. Eye temperature is another promising parameter for assessment of psychological stress, but the technique is likewise susceptible to application errors. Given the high susceptibility of physiological parameters to errors at various experimental stages, behavioural rather than physiological parameters may in fact provide more accurate measures of valence when conducting experiments in the exercising horse. Behavioural parameters that appear to be particularly practical in assessing stress in ridden horses' behaviour are associated with frequencies of behaviour indicative of conflict. However, while increased frequencies of are a good indicator of stress, the absence of conflict behaviour does not provide proof of the absence of stress due to the possible occurrence of conditions such as Learned Helplessness. In future studies, the above issues should be taken into consideration when designing experiments to assess psychological stress in ridden horses.
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Cassinello, J., & Pieters, I. (2000). Multi-male captive groups of endangered dama gazelle: Social rank, aggression, and enclosure effects. Zoo Biol., 19(2), 121–129.
Abstract: A study carried out in four multi-male groups of captive dama gazelle (Gazella dama mhorr) characterized the social rank order of males and possible enclosure effects on aggression rate. A strong relationship between rank and age was found. The results also showed that dominant individuals in the two smallest enclosures were more aggressive than their herdmates, suggesting a more stressful environment, which might precipitate unstable or challenged hierarchies when the animals live in a more restricted enclosure. Subordinate males performed submissive responses at a higher frequency, irrespective of the size of the enclosure. The frequency of interactions between the gazelles, on the other hand, was affected by enclosure size, since high-ranking males showed higher values than low-ranking males in the two smallest enclosures. Frequencies of aggressive acts, retreats, and related interactions were similar in all the herds. Implications for the management of the species in captivity are discussed. Zoo Biol 19:121–129, 2000. © 2000 Wiley-Liss, Inc.
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