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König von Borstel, U., Euent, S., Graf, P., König, S., & Gauly, M. (2011). Equine behaviour and heart rate in temperament tests with or without rider or handler. Physiol. Behav., 104(3), 454–463.
Abstract: The aim of the present study was to compare horses' heart rate (HR), heart rate variability (RMSSD, pNN50) and behaviour in the same temperament test when being ridden, led, and released free. Behavioural measurements included scores and linear measurements for reactivity (R), activity (A), time to calm down (T) and emotionality (E), recorded during the approach (1) and/or during confrontation with the stimulus (2). Sixty-five horses were each confronted 3 times (1 ridden, 1 led, 1 free running in balanced order) with 3 novel and/or sudden stimuli. Mixed model analysis indicated that leading resulted in the lowest (P < 0.05 throughout) reactions as measured by A1, A2, E1, E2, R2, and pNN50 while riding produced the strongest (A1, T2, HR, RMSSD, pNN50) or medium (E1, E2, R2) reactions. Free running resulted either in the strongest (A2, E1, E2, R2) or in the lowest (A1, T2, HR, RMSSD, pNN50) reactions. The repeatability across tests for HR (0.57), but not for RMSSD (0.23) or pNN50 (0.25) was higher than for any behavioural measurement: the latter ranged from values below 0.10 (A1, A2, T2) to values between 0.30 and 0.45 (E1, E2, R2). Overall, the results show that a rider or handler influences, but not completely masks, the horses' intrinsic behaviour in a temperament test, and this influence appeared to be stronger on behavioural variables and heart rate variability than on the horses' heart rates. Taking both practical considerations and repeatabilities into account, reactivity appears to be the most valuable parameter. Emotionality and heart rate can also yield valid results reflecting additional dimensions of temperament although their practical relevance may be less obvious. If a combination of observed variables is chosen with care, a valid assessment of a horse's temperament may be possible in all types of tests. However, in practice, tests that resemble the practical circumstances most closely, i.e. testing riding horses under a rider, should be chosen.
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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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Heffner, R. S., & Heffner, H. E. (1986). Localization of tones by horses: use of binaural cues and the role of the superior olivary complex. Behav Neurosci, 100(1), 93–103.
Abstract: The ability of horses to use binaural time and intensity difference cues to localize sound was assessed in free-field localization tests by using pure tones. The animals were required to discriminate the locus of a single tone pip ranging in frequency from 250 Hz to 25 kHz emitted by loudspeakers located 30 degrees to the left and right of the animals' midline (60 degrees total separation). Three animals were tested with a two-choice procedure; 2 additional animals were tested with a conditioned avoidance procedure. All 5 animals were able to localize 250 Hz, 500 Hz, and 1 kHz but were completely unable to localize 2 kHz and above. Because the frequency of ambiguity for the binaural phase cue delta phi for horses in this test was calculated to be 1.5 kHz, these results indicate that horses can use binaural time differences in the form of delta phi but are unable to use binaural intensity differences. This finding was supported by an unconditioned orientation test involving 4 additional horses, which showed that horses correctly orient to a 500-Hz tone pip but not to an 8-kHz tone pip. Analysis of the superior olivary complex, the brain stem nucleus at which binaural interactions first take place, reveals that the lateral superior olive (LSO) is relatively small in the horse and lacks the laminar arrangement of bipolar cells characteristic of the LSO of most mammals that can use binaural delta I.
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Watanabe, N. M., Stahlman, W. D., Blaisdell, A. P., Garlick, D., Fast, C. D., & Blumstein, D. T. (2012). Quantifying personality in the terrestrial hermit crab: Different measures, different inferences. Behav. Process., 91(2), 133–140.
Abstract: There is much interest in studying animal personalities but considerable debate as to how to define and evaluate them. We assessed the utility of one proposed framework while studying personality in terrestrial hermit crabs (Coenobita clypeatus). We recorded the latency of individuals to emerge from their shells over multiple trials in four unique manipulations. We used the specific testing situations within these manipulations to define two temperament categories (shyness-boldness and exploration-avoidance). Our results identified individual behavioral consistency (i.e., personality) across repeated trials of the same situations, within both categories. Additionally, we found correlations between behaviors across contexts (traits) that suggested that the crabs had behavioral syndromes. While we found some correlations between behaviors that are supposed to measure the same temperament trait, these correlations were not inevitable. Furthermore, a principal component analysis (PCA) of our data revealed new relationships between behaviors and provided the foundation for an alternate interpretation: measured behaviors may be situation-specific, and may not reflect general personality traits at all. These results suggest that more attention must be placed on how we infer personalities from standardized methods, and that we must be careful to not force our data to fit our frameworks.
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Menke, C., Waiblinger, S., Foelsch, D. W., & Wiepkema, P. R. (1999). Social Behaviour and Injuries of Horned Cows in Loose Housing Systems. Anim Welfare, 8(3), 243–258.
Abstract: The relationship between social behaviour and skin injuries (caused by horns) of loose housed horned cows was investigated on 35 dairy farms. While the frequencies of two agonistic behaviour elements (push and chase away) were positively correlated with the occurrence of skin injuries, the frequencies of butting and homing were not. Butting appears to have an ambivalent motivation, in that its occurrence is correlated positively both with agonistic behaviour and with social licking. Horning showed a positive correlation with social licking only. Four groups of husbandry conditions that may be associated with the occurrence of social behaviour and of injuries were distinguished: i) herd management, with variables including problem solving management by the farmer, integration of new cows, and dealing with periparturient and oestrus cows; ii) human-animal relationship, with variables including ability to identify individual cows, frequency of brushing the cows, number of milkers, and frequency of personnel changes; iii) animal characteristics, with the variable of herd size; and iv) stable characteristics, with the variable of space per cow (m2). The relevance of the husbandry variables investigated here had been confirmed in a previous stepwise regression analysis (Menke 1996). The variables for herd management and human-animal relationship conditions correlated in a consistent way with the occurrence of agonistic behaviour and/or of injuries, while most of them also correlated in the opposite direction with the occurrence of social licking. Herd size correlated positively with agonistic behaviour, but negatively with social licking. Space per cow correlated negatively with agonistic behaviour and injuries. In more than 70 per cent of the herds investigated, the levels of agonistic behaviour and of skin injuries were low, implying that horned dairy cows can be kept with less risk than is often assumed. We argue that such risks strongly depend on management factors that can be improved.
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Li, C., Jiang, Z., Tang, S., & Zeng, Y. (2007). Influence of enclosure size and animal density on fecal cortisol concentration and aggression in Pere David's deer stags. Gen Comp Endocrinol, 151(2), 202–209.
Abstract: We investigated the impact of enclosure size and animal density on behavior and adrenocortical secretion in Pere David's deer in Dafeng Nature Reserve, China. From February 15 to April 16 in 2004, we conducted two experiments. First, we studied maintenance behavior and conflict behavior of Pere David's deer stags in a large enclosure (200 ha) with low animal density (0.66 deer/ha) and a small display pen (0.75 ha) with high animal density (25.33 deer/ha). The maintenance behavior we recorded included standing, locomotion, foraging and rest. During the behavioral observations, we collected fresh voided fecal samples from the stags periodically, and analyzed the fecal cortisol concentrations in those samples using radioimmunoassay technique. Second, we monitored the fecal cortisol concentrations of one group of stags (12 deer lived in an enclosure of 100 ha) before and after transferred into a small pen (0.5 ha). We found that in the first experiment: (1) there were significant differences in standing and rest whereas no significant differences of locomotion and foraging between the free-ranging group and the display group; (2) frequency of conflict behavior in the display group was significantly higher than those in the free-ranging group; and (3) fecal cortisol concentration of the display group (326.17+/-16.98 ng/g dry feces) was significantly higher than that of the free-ranging group (268.98+/-15.21 ng/g dry feces). In the second experiment, there was no significant difference of the fecal cortisol concentrations among sampling days, but the mean fecal cortisol concentration of the day after transferring (337.46+/-17.88 ng/g dry feces) was significantly higher than that of the day before transferring (248.44+/-7.99 ng/g dry feces). Comparison with published findings, our results indicated that enclosure size and animal density affect not only behaviors, but also adrenocortical secretion in Pere David's deer. Small living space with high animal density may impose physiological stress to captive Pere David's deer. Moreover, long-term physiological stress and increase of conflict behavior may subsequently affect survival and reproduction of the deer.
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Lee, R. D. (2003). Rethinking the evolutionary theory of aging: transfers, not births, shape senescence in social species. Proc Natl Acad Sci U S A, 100(16), 9637–9642.
Abstract: The classic evolutionary theory of aging explains why mortality rises with age: as individuals grow older, less lifetime fertility remains, so continued survival contributes less to reproductive fitness. However, successful reproduction often involves intergenerational transfers as well as fertility. In the formal theory offered here, age-specific selective pressure on mortality depends on a weighted average of remaining fertility (the classic effect) and remaining intergenerational transfers to be made to others. For species at the optimal quantity-investment tradeoff for offspring, only the transfer effect shapes mortality, explaining postreproductive survival and why juvenile mortality declines with age. It also explains the evolution of lower fertility, longer life, and increased investments in offspring.
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Krueger, K., & Farmer, K. (2011). Laterality in the Horse [Lateralität beim Pferd ]. mup, 4, 160–167.
Abstract: Horses are one-sided, not only on a motor level, but they also prefer to use one eye, ear or nostril over the other under particular circumstances. Horses usually prefer using the left eye to observe novel objects and humans. This preference is more marked in emotional situations and when confronted with unknown persons. Thus the horse’s visual laterality provides a good option for assessing its mental state during training or in human-horse interactions. A strong preference for the left eye may signal that a horse cannot deal with certain training situations or is emotionally affected by a particular person.
Pferde benutzen für die Begutachtung von Objekten und Menschen bevorzugt eine bestimmte Nüster, ein Ohr oder ein Auge. So betrachten die meisten Pferde Objekte und Menschen mit dem linken Auge. Die Lateralitätsforschung erklärt diese sensorische Lateralität mit der Verarbeitung von Informationen unterschiedlicher Qualität in verschiedenen Gehirnhälften und zeigt auf, dass positive und negative emotionale Informationen sowie soziale Sachverhalte mit dem linken Auge aufgenommen und vorwiegend an die rechte Gehirnhälfte weitergegeben werden. In diesem Zusammenhang ermöglicht die visuelle Lateralität, den Gemütszustand des Pferdes im Training und im therapeutischen Fördereinsatz zu erkennen und zu berücksichtigen.
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Sankey, C., Richard-Yris, M. - A., Leroy, H., Henry, S., & Hausberger, M. (2010). Positive interactions lead to lasting positive memories in horses, Equus caballus. Anim. Behav., 79(4), 869–875.
Abstract: Social relationships are important in social species. These relationships, based on repeated interactions, define each partner's expectations during the following encounters. The creation of a relationship implies high social cognitive abilities which require that each partner is able to associate the positive or negative content of an interaction with a specific partner and to recall this association. In this study, we tested the effects of repeated interactions on the memory kept by 23 young horses about humans, after 6 and 8 months of separation. The association of a reward with a learning task in an interactional context induced positive reactions towards humans during training. It also increased contact and interest, not only just after training, but also several months later, despite no further interaction with humans. In addition, this ‘positive memory’ of humans extended to novel persons. Overall, positive reinforcement enhanced learning and memorization of the task itself. These findings suggest remarkable social cognitive abilities that can be transposed from intraspecific to interspecific social contexts.
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Weissing, F. J. (2011). Animal behaviour: Born leaders. Nature, 474(7351), 288–289.
Abstract: Social animals face a dilemma. To reap the benefits of group living, they have to stay together. However, individuals differ in their preferences as to where to go and what to do next. If all individuals follow their own preferences, group coherence is undermined, resulting in an outcome that is unfavourable for everyone. Neglecting one's own preferences and following a leader is one way to resolve this coordination problem. But what attributes make an individual a 'leader'? A modelling study by Johnstone and Manica1 illuminates this question.
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