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Massen, J., Sterck, E., & de Vos, H. (2010). Close social associations in animals and humans: functions and mechanisms of friendship (Vol. 147).
Abstract: 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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Minero, M., Canali, E., Ferrante, V., Verga, M., & Odberg, F. O. (1999). Heart rate and behavioural responses of crib-biting horses to two acute stressors. Vet. Rec., 145(15), 430–433.
Abstract: The heart rate and behaviour of 14 adult saddle horses, eight crib-biters and six normal controls, were investigated. Initially, the relationship between crib-biting and heart rate was investigated while the horses were undisturbed. The horses were tested when restrained with a lip twitch, and assessed when they were exposed suddenly to the rapid inflation of a balloon. The heart rate of the crib-biters during crib-biting was lower than during other behaviours. The crib-biters had a higher overall mean heart rate (P<0.05) suggesting that they may have had a higher basal sympathetic activity. After the application of the twitch, all the horses had a transient increase in heart rate which returned to basal values more rapidly in the crib-biters. The crib-biters were less reactive to the lip twitch, five of the six investigated remaining calm, and after the release of the twitch, they spent more time nibbling (P<0.05) than the control horses. The crib-biters reacted more strongly to the inflation of the balloon (three of the six reacted), and after it had been inflated they spent more time walking in the box.
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Christensen, J. W., Ahrendt, L. P., Lintrup, R., Gaillard, C., Palme, R., & Malmkvist, J. (2012). Does learning performance in horses relate to fearfulness, baseline stress hormone, and social rank? In Applied Animal Behaviour Science (Vol. 140, pp. 44–52).
Abstract: 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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Ganswindt, A., Palme, R., Heistermann, M., Borragan, S., & Hodges, J. K. (2003). Non-invasive assessment of adrenocortical function in the male African elephant (Loxodonta africana) and its relation to musth. Gen Comp Endocrinol, 134(2), 156–166.
Abstract: Adult male elephants periodically show the phenomenon of musth, a condition associated with increased aggressiveness, restlessness, significant weight reduction and markedly elevated androgen levels. It has been suggested that musth-related behaviours are costly and that therefore musth may represent a form of physiological stress. In order to provide data on this largely unanswered question, the first aim of this study was to evaluate different assays for non-invasive assessment of adrenocortical function in the male African elephant by (i) characterizing the metabolism and excretion of [3H]cortisol (3H-C) and [14C]testosterone (14C-T) and (ii) using this information to evaluate the specificity of four antibodies for determination of excreted cortisol metabolites, particularly with respect to possible cross-reactions with androgen metabolites, and to assess their biological validity using an ACTH challenge test. Based on the methodology established, the second objective was to provide data on fecal cortisol metabolite concentrations in bulls during the musth and non-musth condition. 3H-C (1 mCi) and 14C-T (100 microCi) were injected simultaneously into a 16 year old male and all urine and feces collected for 30 and 86 h, respectively. The majority (82%) of cortisol metabolites was excreted into the urine, whereas testosterone metabolites were mainly (57%) excreted into the feces. Almost all radioactive metabolites recovered from urine were conjugated (86% 3H-C and 97% 14C-T). In contrast, 86% and >99% of the 3H-C and 14C-T metabolites recovered from feces consisted of unconjugated forms. HPLC separations indicated the presence of various metabolites of cortisol in both urine and feces, with cortisol being abundant in hydrolysed urine, but virtually absent in feces. Although all antibodies measured substantial amounts of immunoreactivity after HPLC separation of peak radioactive samples and detected an increase in glucocorticoid output following the ACTH challenge, only two (in feces against 3alpha,11-oxo-cortisol metabolites, measured by an 11-oxo-etiocholanolone-EIA and in urine against cortisol, measured by a cortisol-EIA) did not show substantial cross-reactivity with excreted 14C-T metabolites and could provide an acceptable degree of specificity for reliable assessment of glucocorticoid output from urine and feces. Based on these findings, concentrations of immunoreactive 3alpha,11-oxo-cortisol metabolites were determined in weekly fecal samples collected from four adult bulls over periods of 11-20 months to examine whether musth is associated with increased adrenal activity. Results showed that in each male levels of these cortisol metabolites were not elevated during periods of musth, suggesting that in the African elephant musth is generally not associated with marked elevations in glucocorticoid output. Given the complex nature of musth and the variety of factors that are likely to influence its manifestation, it is clear, however, that further studies, particularly on free-ranging animals, are needed before a possible relationship between musth and adrenal function can be resolved. This study also clearly illustrates the potential problems associated with cross-reacting metabolites of gonadal steroids in EIAs measuring glucocorticoid metabolites. This has to be taken into account when selecting assays and interpreting results of glucocorticoid metabolite analysis, not only for studies in the elephant but also in other species.
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Oikawa, M., Hobo, S., Oyamada, T., & Yoshikawa, H. (2005). Effects of Orientation, Intermittent Rest and Vehicle Cleaning During Transport on Development of Transport-related Respiratory Disease in Horses. Journal of Comparative Pathology, 132(2-3), 153–168.
Abstract: Summary The effects of various factors on the inflammatory and stress response in horses during transportation were examined in Experiments 1 and 2, carried out in April and August, respectively. In Experiment 1, three groups (G1-G3) of four Thoroughbreds were used, and in Experiment 2, two groups (G4, G5). G1 animals were loaded into lorries with their heads facing forwards (FF) and given periods of short rest (SR) (30 min for every 4 h driven). G2 horses were loaded facing rearwards (FR) and given SR. G3 horses were FF and given periods of long rest (LR) (2 h rest for every 4 h driven). G4 horses were transported FF with hay suspended in front of them and no cleaning of faeces or urine during rest stops. G5 horses were FF and given pellets instead of hay, and the lorry was cleaned and washed at each rest stop. G4 and G5 horses were rested for 1 h after every 5 h of driving. All groups were driven a distance of 1500 km, and the total journey time was 37 h for G1 and G2, 49 h for G3, and 40 h for G4 and G5. In Experiment 1, indicators of inflammatory and stress responses tended to be highest in G1, intermediate in G2 and lowest in G3. In Experiment 2 they tended to be higher in G4 than in G5. The results suggested that increasing the rest time and cleaning the interior of the vehicle during rest stops reduced transportation stress and respiratory insults, factors that may lead to respiratory disease.
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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., et al. (2006). 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. Tijdschr Diergeneeskd, 131(5), 152–157.
Abstract: '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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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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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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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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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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