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Touma, C., Sachser, N., Mostl, E., & Palme, R. (2003). Effects of sex and time of day on metabolism and excretion of corticosterone in urine and feces of mice. Gen Comp Endocrinol, 130(3), 267–278.
Abstract: Non-invasive techniques to monitor stress hormones in small animals like mice offer several advantages and are highly demanded in laboratory as well as in field research. Since knowledge about the species-specific metabolism and excretion of glucocorticoids is essential to develop such a technique, we conducted radiometabolism studies in mice (Mus musculus f. domesticus, strain C57BL/6J). Each mouse was injected intraperitoneally with 740 kBq of 3H-labelled corticosterone and all voided urine and fecal samples were collected for five days. In a first experiment 16 animals (eight of each sex) received the injection at 9 a.m., while eight mice (four of each sex) were injected at 9 p.m. in a second experiment. In both experiments radioactive metabolites were recovered predominantly in the feces, although males excreted significantly higher proportions via the feces (about 73%) than females (about 53%). Peak radioactivity in the urine was detected within about 2h after injection, while in the feces peak concentrations were observed later (depending on the time of injection: about 10h postinjection in experiment 1 and about 4h postinjection in experiment 2, thus proving an effect of the time of day). The number and relative abundance of fecal [3H]corticosterone metabolites was determined by high performance liquid chromatography (HPLC). The HPLC separations revealed that corticosterone was extensively metabolized mainly to more polar substances. Regarding the types of metabolites formed, significant differences were found between males and females, but not between the experiments. Additionally, the immunoreactivity of these metabolites was assessed by screening the HPLC fractions with four enzyme immunoassays (EIA). However, only a newly established EIA for 5alpha-pregnane-3beta,11beta,21-triol-20-one (measuring corticosterone metabolites with a 5alpha-3beta,11beta-diol structure) detected several peaks of radioactive metabolites with high intensity in both sexes, while the other EIAs showed only minor immunoreactivity. Thus, our study for the first time provides substantial information about metabolism and excretion of corticosterone in urine and feces of mice and is the first demonstrating a significant impact of the animals' sex and the time of day. Based on these data it should be possible to monitor adrenocortical activity non-invasively in this species by measuring fecal corticosterone metabolites with the newly developed EIA. Since mice are extensively used in research world-wide, this could open new perspectives in various fields from ecology to behavioral endocrinology.
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Romero L. M. (2011). Using the reactive scope model to understand why stress physiology predicts survival during starvation in Galápagos marine iguanas. Gen Comp Endocrinol, .
Abstract: 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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Touma, C., Palme, R., & Sachser, N. (2004). Analyzing corticosterone metabolites in fecal samples of mice: a noninvasive technique to monitor stress hormones. Horm Behav, 45(1), 10–22.
Abstract: In small animals like mice, the monitoring of endocrine functions over time is constrained seriously by the adverse effects of blood sampling. Therefore, noninvasive techniques to monitor, for example, stress hormones in these animals are highly demanded in laboratory as well as in field research. The aim of our study was to evaluate the biological relevance of a recently developed technique to monitor stress hormone metabolites in fecal samples of laboratory mice. In total, six experiments were performed using six male and six female mice each. Two adrenocorticotropic hormone (ACTH) challenge tests, two dexamethasone (Dex) suppression tests and two control experiments [investigating effects of the injection procedure itself and the diurnal variation (DV) of glucocorticoids (GCs), respectively] were conducted. The experiments clearly demonstrated that pharmacological stimulation and suppression of adrenocortical activity was reflected accurately by means of corticosterone metabolite (CM) measurements in the feces of males and females. Furthermore, the technique proved sensitive enough to detect dosage-dependent effects of the ACTH/Dex treatment and facilitated to reveal profound effects of the injection procedure itself. Even the naturally occurring DV of GCs could be monitored reliably. Thus, our results confirm that measurement of fecal CM with the recently established 5alpha-pregnane-3beta,11beta,21-triol-20-one enzyme immunoassay is a very powerful tool to monitor adrenocortical activity in laboratory mice. Since mice represent the vast majority of all rodents used for research worldwide and the number of transgenic and knockout mice utilized as animal models is still increasing, this noninvasive technique can open new perspectives in biomedical and behavioral science.
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Reimers, M., Schwarzenberger, F., & Preuschoft, S. (2007). Rehabilitation of research chimpanzees: stress and coping after long-term isolation. Horm Behav, 51(3), 428–435.
Abstract: We report on the permanent retirement of chimpanzees from biomedical research and on resocialization after long-term social isolation. Our aim was to investigate to what extent behavioral and endocrine measures of stress in deprived laboratory chimpanzees can be improved by a more species-typical social life style. Personality in terms of novelty responses, social dominance after resocialization and hormonal stress susceptibility were affected by the onset of maternal separation of infant chimpanzees and duration of deprivation. Chimpanzees, who were separated from their mothers at a younger age and kept in isolation for more years appeared to be more timid personalities, less socially active, less dominant and more susceptible to stress, as compared to chimpanzees with a less severe deprivation history. However, permanent retirement from biomedical research in combination with therapeutic resocialization maximizing chimpanzees' situation control resulted in reduced fecal cortisol metabolite levels. Our results indicate that chimpanzees can recover from severe social deprivation, and may experience resocialization as less stressful than solitary housing.
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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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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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Dreschel, N. A., & Granger, D. A. (2009). Methods of collection for salivary cortisol measurement in dogs. Horm. Behav., 55(1), 163–168.
Abstract: 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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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.
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Alexander, F., & Ash, R. W. (1955). The effect of emotion and hormones on the concentration of glucose and eosinophils in horse blood. J Physiol, 130(3), 703–710.
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Hausberger, M., Fureix, C., Bourjade, M., Wessel-Robert, S., & Richard-Yris, M. - A. (2012). On the significance of adult play: what does social play tell us about adult horse welfare? Naturwissenschaften, 99(4), 291–302.
Abstract: 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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