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Call, J., Aureli, F., & de Waal, F. B. M. (2002). Postconflict third-party affiliation in stumptailed macaques. Anim. Behav., 63(2), 209–216.
Abstract: Stumptailed macaques, Macaca arctoides, are characterized by high levels of postconflict affiliative contacts between opponents. We investigated the occurrence of postconflict affiliative contacts between opponents and third parties that were not involved in the original conflict. We collected 10-min focal observations during postconflict and control periods in which we recorded all aggressive and affiliative behaviours between opponents and third parties. We distinguished three types of third parties depending on the relationship with the focal animal: own kin, opponent's kin and individuals unrelated to both opponents. We analysed the interactions with third parties separately, while distinguishing two classes of affiliative behaviours: (1) allogrooming and contact sitting and (2) sociosexual behaviours (e.g. genital inspection). The macaques showed differences between postconflict and control periods in their affiliative contacts with third parties. Aggressors received more postconflict grooming and contact sitting from their opponents' kin, received more sociosexual behaviour from their own kin and unrelated individuals, and directed more sociosexual behaviour to unrelated individuals. Victims received and directed less postconflict grooming from and towards their own kin. They received more postconflict sociosexual behaviour from all partners except their own kin and directed more sociosexual behaviour to all partners except the opponent's kin. This study establishes the occurrence of multiple postconflict triadic affiliation in stumptailed macaques, and is the first to show that victims receive contacts from third parties in a cercopithecine species, a behaviour previously described only in chimpanzees. It also highlights the importance of analysing the different affiliative behaviours separately in postconflict situations. Otherwise, many of the patterns we report, especially those involving victims, would have been missed.
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Koski, S. E., & Sterck, E. H. M. (2007). Triadic postconflict affiliation in captive chimpanzees: does consolation console? Anim. Behav., 73(1), 133–142.
Abstract: Consolation is a triadic postconflict interaction between a conflict participant and an uninvolved third party. The term consolation implies stress alleviation. Consequently, consolation may be an effective mechanism to alleviate postconflict stress. However, this assumption has not been tested. We tested whether consolation alleviates postconflict stress in captive chimpanzees, Pan troglodytes. In addition, we examined whether consolation is a substitute postconflict interaction for reconciliation. We collected 643 postconflict-matched control pairs on aggressees and 576 on aggressors. Consolation occurred equally frequently with aggressees and aggressors. However, we found no evidence that consolation alleviated stress, regardless of the identity of the consoler. In addition, consolation was also directed to conflict participants with no evident postconflict stress. Furthermore, we found no evidence for consolation being a substitute for reconciliation. The occurrence of consolation did not depend on the occurrence of reconciliation and consolation was not more prevalent with the sex class that reconciled less often or had the highest postconflict stress levels. We conclude that consolation is a postconflict interaction in its own right, the function of which is not likely to be connected to stress alleviation of the consoled individual. We propose that the function of triadic postconflict affiliation, previously labelled as consolation, should be reassessed with regard to the third parties' reasons to affiliate with conflict opponents.
Keywords: chimpanzee; consolation; Pan troglodytes; postconflict affiliation
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Mills, D. S., & Riezebos, M. (2005). The role of the image of a conspecific in the regulation of stereotypic head movements in the horse. Appl. Anim. Behav. Sci., 91(1-2), 155–165.
Abstract: Weaving is a common locomotor stereotypy in horses, which has been shown to be reduced by the presence of a mirror in the stable; however, it remains unclear whether this effect is due to the reflected image of a horse or some other property associated with the introduction of a mirror. To investigate this further, the response of six known weaving horses to each of three different types of poster was recorded. The poster images consisted of a life-size image of a horse's face (true image), a version of the same image cut into 54 squares, which were then randomly rearranged (pixilated image) and a blank (white image) display of the same size as the other posters. Four observation periods were included in each of the two days of observation per treatment condition: 08:00-08:30 h, 10:00-10:30 h, 12:00-12:30 h and 16:00-17:00 h. The first observation period was followed by the provision of concentrate feed and forage, the second observation period by exercise, the third by further forage and the fourth by concentrate feed and forage. The horses' activities and positions in the box were scanned at 1 min intervals and the number of observations registering each activity and position was calculated as a percentage of the total number of observations for each observation period. Weaving was significantly less when the horses were provided with the image of a horse's face (mean percentage of observations +/- S.E.M.; 5.56 +/- 1.57), compared to both the pixilated (14.85 +/- 3.06) and white (20.52 +/- 4.12) images. Nodding was significantly less when provided with the true image of a horse (0.45 +/- 0.15) compared to the pixilated image (2.15 +/- 0.67). The pattern of weaving throughout the day differed from that of nodding, and together these results suggest that the two behaviours are not analogous. The presence of the image of the horse's face was associated with a significant increase in alertness (F112,2 = 11.31, p < 0.001) and recorded time spent looking at the poster(F112,2 = 3.46, p < 0.05). The timing of stereotypic head movements in the horse in this study suggests that they are not associated with boredom, or lack of stimulation but rather acute frustration.
Keywords: Behaviour; Horse; Nodding; Perception; Stereotypy; Vision; Weaving
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Sondergaard, E., & Halekoh, U. (2003). Young horses' reactions to humans in relation to handling and social environment. Appl. Anim. Behav. Sci., 84(4), 265–280.
Abstract: Forty Danish warmblood colts in two replicates were used to investigate the effect of housing and handling in the rearing period on the reactions to humans. The horses entered the experiment after weaning and were housed either individually (n=16) or in groups of three (n=24). Half of the horses from each housing group were handled three times per week for a period of 10 min. Approach tests were performed in the home environment when the horses were 6, 9, 12, 18, 21, and 24 months old, and an Arena and Human Encounter test was performed in a novel environment when the horses were 12 and 24 months old, respectively. In the home environment, single-housed horses approached sooner and were more easily approached by a human than group-housed horses where no effect of handling was observed. Horses approached sooner and were more easily approached with increasing age. In the Arena and Human Encounter test, single-housed horses expressed less restless behaviour, more explorative behaviour, and less vocalisation than group-housed horses. Handled horses showed lower increase in heart rate during the test than non-handled horses. There was no difference between the number of times single or group-housed horses touched an unfamiliar person in the Arena and Human Encounter test but handled horses approached sooner than non-handled horses. It is concluded that the social environment affected the way horses reacted to humans when tested in the home environment but not in a novel environment. In contrast, handling affected the reactions to humans when tested in the novel environment but not in the home environment. However, handled horses also reacted less to the novel environment in general, thus indicating that handling is a mean of avoiding potential dangerous situations.
Keywords: Horses; Human-animal relationship; Social environment; Handling
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Rietmann, T. R., Stuart, A. E. A., Bernasconi, P., Stauffacher, M., Auer, J. A., & Weishaupt, M. A. (2004). Assessment of mental stress in warmblood horses: heart rate variability in comparison to heart rate and selected behavioural parameters. Appl. Anim. Behav. Sci., 88(1-2), 121–136.
Abstract: The aim of the study was to investigate whether heart rate variability (HRV) could assess alterations of the autonomic nervous system (ANS) at different levels of excitement. The behavioural and physiological responses of 20 warmblood horses to a challenging ground exercise task were studied. Prior to the experiment, the horses were evaluated at rest and during forward walking (FW). The horses were then forced to move backwards continuously during 3 min according to a standardised protocol (BW1). Subsequently, the horses were exposed to two training sessions, after which the backward walking (BW2) was re-evaluated. Heart rate (HR) and HRV-parameters such as the standard deviation of the beat-to-beat intervals (SDRR), the low (LF; sympathetic tone) and high frequency (HF) component of HRV (HF; parasympathetic tone) and their ratio (LF/HF; index representing the sympatho-vagal balance) were sampled at rest, and during FW, BW1 and BW2. Stress-related behaviour during BW1 and BW2 was determined from video recordings. The results of the different evaluations were compared to each other. Compared to rest and FW, the first backward experiment induced a significant rise in HR, LF and LF/HF and a significant decrease of HF. SDRR decreased from both FW and rest with only the latter reaching significance. In BW2 after the training sessions, HR and the parameters of the sympathetic branch of the ANS (LF, LF/HF) were decreased and the vagal tone (HF) increased compared to BW1; all changes were significant. The duration of stress indicating behavioural patterns revealed also a significant decrease of excitement after the training, when backward walking did not differ from forward walking in any parameter. Correlations between HRV-parameters and stress indicating behaviour as well as HR were found. We conclude that the HRV-parameters LF and HF are valuable measures to quantify sympatho-vagal balance, which allows a more precise assessment of the responses of HR and SDRR to mental stress during low intensity exercise.
Keywords: Horse; Hrv; Heart rate; Mental stress; Behaviour; Autonomic nervous system
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Sibbald, A. M., Elston, D. A., Smith, D. J. F., & Erhard, H. W. (2005). A method for assessing the relative sociability of individuals within groups: an example with grazing sheep. Appl. Anim. Behav. Sci., 91(1-2), 57–73.
Abstract: We describe a method for quantifying relative sociability within a group of animals, which is defined as the tendency to be close to others within the group and based on the identification of nearest neighbours. The method is suitable for groups of animals in which all individuals are visible and identifiable and has application as a tool in other areas of behavioural research. A sociability index (SI) is calculated, which is equivalent to the relative proportion of time that an individual spends as the nearest neighbour of other animals in the group and is scaled to have an expectation of 1.0 under the null hypothesis of random mixing. Associated pairs, which are animals seen as nearest neighbours more often than would be expected by chance, are also identified. The method tests for consistency across a number of independent observation periods, by comparison with values obtained from simulations in which animal identities are randomised between observation periods. An experiment is described in which 8 groups of 7 grazing sheep were each observed for a total of 10, one-hour periods and the identities and distances away of the 3 nearest neighbours of each focal animal recorded at 5-min intervals. Significant within-group differences in SIs were found in four of the groups (P < 0.001). SIs calculated using the nearest neighbour, two nearest neighbours or three nearest neighbours, were generally highly correlated within all groups, with little change in the ranking of animals. There were significant negative correlations between SIs and nearest neighbour distances in five of the groups. It was concluded that there was no advantage in recording more than one neighbour to calculate the SI. Advantages of the SI over other methods for measuring sociability and pair-wise associations are discussed.
Keywords: Association; Grazing; Nearest neighbour; Sheep; Sociability; Social behaviour
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Heitor, F., do Mar Oom, M., & Vicente, L. (2006). Social relationships in a herd of Sorraia horses: Part II. Factors affecting affiliative relationships and sexual behaviours. Behav. Process., 73(3), 231–239.
Abstract: The influence of age, dominance rank, kinship and aggressiveness over affiliative relationships and sexual behaviours were analysed in a herd of Sorraia horses, Equus caballus, kept under extensive management. Subjects were 10 adult mares 5-18 years old that had known each other since birth, and a stallion introduced into the group for breeding for the first time. Kinship coefficient and dominance rank were the most important factors affecting affiliative relationships. Bonds were reciprocal and stronger among mares with higher kinship. Mares spent more time in proximity to close-ranking and lower-ranking females. Mares with stronger affiliative relationships or higher relatedness were not less aggressive towards each other. Affiliative relationships between the stallion and the mares were not reciprocal: lower-ranking mares formed stronger bonds with the stallion but he preferred the less genetically related mares for proximity. However, the stallion was involved in sexual behaviours more frequently with the mares that were more genetically related to him. These results suggest that kinship beyond close relatives may affect affiliative relationships both among familiar and among unfamiliar horses. However, the influence of kinship does not imply that horses possess a kin recognition system and alternative explanations are discussed.
Keywords: Horse; Equus; Affiliative relationship; Rank; Kinship; Aggressiveness
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Visser, E. K., van Reenen, C. G., van der Werf, J. T. N., Schilder, M. B. H., Knaap, J. H., Barneveld, A., et al. (2002). Heart rate and heart rate variability during a novel object test and a handling test in young horses. Physiol. Behav., 76(2), 289–296.
Abstract: Forty-one Dutch Warmblood immature horses were used in a study to quantify temperamental traits on the basis of heart rate (HR) and heart rate variability (HRV) measures. Half of the horses received additional training from the age of 5 months onwards; the other half did not. Horses were tested at 9, 10, 21 and 22 months of age in a novel object and a handling test. During the tests, mean HR and two heart variability indices, e.g. standard deviation of beat-to-beat intervals (SDRR) and root mean square of successive beat-to-beat differences (rMSSD), were calculated and expressed as response values to baseline measures. In both tests, horses showed at all ages a significant increase in mean HR and decrease in HRV measures, which suggests a marked shift of the balance of the autonomic nervous system towards a sympathetic dominance. In the novel object test, this shift was more pronounced in horses that had not been trained. Furthermore, statistical analysis showed that the increase in mean HR could not be entirely explained by the physical activity. The additional increase in HR, the nonmotor HR, was more pronounced in the untrained horses compared to the trained. Hence, it is suggested that this nonmotor HR might be due to the level of emotionality. HR variables showed consistency between years, as well as within the second year. These tests bring about a HR response in horses, part of which may indicate a higher level of emotionality; and horses show individual consistency of these HR variables over ages. Therefore, it is concluded that mean HR and HRV measures used with these tests quantify certain aspects of a horse's temperament.
Keywords: Heart rate; Heart rate variability; Consistency; Behavioral tests; Temperament; Emotionality; Horses
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Visser, E. K., van Reenen, C. G., Hopster, H., Schilder, M. B. H., Knaap, J. H., Barneveld, A., et al. (2001). Quantifying aspects of young horses' temperament: consistency of behavioural variables. Appl. Anim. Behav. Sci., 74(4), 241–258.
Abstract: Performance of horses, whether in sports or in leisure, depends on both physical abilities as well as temperament. The aim of the present work was to measure individual variation and consistency of behavioural variables, related to temperament, in young horses of the same breed and age, and reared under controlled housing conditions and management. A total of 41 Dutch Warmblood horses were tested at 9, 10, 21 and 22 months of age in two behavioural tests, i.e. the novel object test and the handling test. In the novel object test horses were confronted with an open umbrella that was lowered from the ceiling. In the handling test horses were led by a human to cross a bridge. Per test, behavioural variables in the following behavioural classes were observed: locomotor activity, latency times, postural expressions and vocalisations. Within years, all behavioural variables in the handling test, and all but two in the novel object test were positively correlated (0.36<Rs<0.81, P<0.05). For both tests, at 9, 10, 21 and 22 months of age, a principal component analysis (PCA) was carried out to examine whether there were indications for underlying components of these individual behavioural variables that could possibly serve as measures for temperamental traits. The first component in the novel object test could be regarded as `flightiness' and the second as `sensitiveness'. In the handling test, the first component was suggested to relate to `patience', the second component to `willingness to perform'. The temperamental trait `flightiness' (novel object test) as well as the temperamental trait `patience' (handling test) were positively correlated within both years (0.36<Rs<0.65, P<0.05). For the traits `sensitiveness' (novel object test) and `willingness to perform' (handling test) a positive correlation was only found within the first year (0.44<Rs<0.57, P<0.01). A few individual behavioural variables showed consistency over years. Additionally, just one out of four temperamental traits, namely `flightiness', proved to be consistent over years (Rs=0.49, P<0.01). The temperamental trait `patience' showed a trend between years (Rs=0.31, 0.05<P<0.1). It is concluded that the behavioural tests employed in the present study can be used to reliably identify individual behavioural variables and temperamental traits in young horses. Long-term consistency, i.e. between subsequent years, could not be demonstrated convincingly. Nevertheless, future work may indicate that employing the same approach and considering an even longer time period or different phases of the horse's life, long-term consistency does exist.
Keywords: Horses; Temperament; Individual differences; Behavioural variables; Pca
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Visser, E. K., Van Reenen, C. G., Engel, B., Schilder, M. B. H., Barneveld, A., & Blokhuis, H. J. (2003). The association between performance in show-jumping and personality traits earlier in life. Appl. Anim. Behav. Sci., 82(4), 279–295.
Abstract: For a horse to succeed in a show-jumping career, the individual has to possess both excellent physical abilities as well as a suitable personality to perform under challenging conditions. Forty-one Dutch Warmblood horses were used to develop personality tests and correlations between test variables and early training performances in jumping were studied. In behavioural tests, during the first 2 years of the horses' lives, personality aspects like emotionality, reactivity to human and learning abilities were quantified. At the age of 3, horses were broken and received early training in show-jumping. The inter-relationship between several performance variables measured during this early training phase were studied using principal component analysis (PCA). Variables measured in the different personality tests (novel-object test, handling test, avoidance-learning test and a reward-learning test) showed no correlations, suggesting that these tests all triggered different aspects of a horse's personality. This study indicates that it is possible to predict a substantial part of the show-jumping performance of an individual horse later in life by personality traits earlier in life.
Keywords: Personality; Performance; Horses; Prediction; Individual differences; Behavioural tests
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