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Mizuguchi, M.; Arai, M.; Ke, Y.; Nitta, K.; Kuwajima, K. |
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Title |
Equilibrium and kinetics of the folding of equine lysozyme studied by circular dichroism spectroscopy |
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Journal Article |
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1998 |
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Journal of Molecular Biology |
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283 |
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1 |
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265-277 |
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equine lysozyme; protein folding; molten globule; stopped-flow; folding intermediate |
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The equilibrium unfolding and the kinetics of unfolding and refolding of equine lysozyme, a Ca2+-binding protein, were studied by means of circular dichroism spectra in the far and near-ultraviolet regions. The transition curves of the guanidine hydrochloride-induced unfolding measured at 230 nm and 292.5 nm, and for the apo and holo forms of the protein have shown that the unfolding is well represented by a three-state mechanism in which the molten globule state is populated as a stable intermediate. The molten globule state of this protein is more stable and more native-like than that of α-lactalbumin, a homologous protein of equine lysozyme. The kinetic unfolding and refolding of the protein were induced by concentration jumps of the denaturant and measured by stopped-flow circular dichroism. The observed unfolding and refolding curves both agreed well with a single-exponential function. However, in the kinetic refolding reactions below 3 M guanidine hydrochloride, a burst-phase change in the circular dichroism was present, and the burst-phase intermediate in the kinetic refolding is shown to be identical with the molten globule state observed in the equilibrium unfolding. Under a strongly native condition, virtually all the molecules of equine lysozyme transform the structure from the unfolded state into the molten globule, and the subsequent refolding takes place from the molten globule state. The transition state of folding, which may exist between the molten globule and the native states, was characterized by investigating the guanidine hydrochloride concentration-dependence of the rate constants of refolding and unfolding. More than 80% of the hydrophobic surface of the protein is buried in the transition state, so that it is much closer to the native state than to the molten globule in which only 36% of the surface is buried in the interior of the molecule. It is concluded that all the present results are best explained by a sequential model of protein folding, in which the molten globule state is an obligatory folding intermediate on the pathway of folding. |
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refbase @ user @ |
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3990 |
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Palm, A.-K.E.; Wattle, O.; Lundström, T.; Wattrang, E. |
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Title |
Secretory immunoglobulin A and immunoglobulin G in horse saliva |
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Journal Article |
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Year |
2016 |
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Veterinary Immunology and Immunopathology |
Abbreviated Journal |
Vet. Immunol. Immunolpathol. |
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180 |
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59-65 |
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Equine; Secretory IgA; IgG; Saliva; Mucosal immunity |
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This study aimed to increase the knowledge on salivary antibodies in the horse since these constitute an important part of the immune defence of the oral cavity. For that purpose assays to detect horse immunoglobulin A (IgA) including secretory IgA (SIgA) were set up and the molecular weights of different components of the horse IgA system were estimated. Moreover, samples from 51 clinically healthy horses were tested for total SIgA and IgG amounts in saliva and relative IgG3/5 (IgG(T)) and IgG4/7 (IgGb) content were tested in serum and saliva. Results showed a mean concentration of 74μg SIgA/ml horse saliva and that there was a large inter-individual variation in salivary SIgA concentration. For total IgG the mean concentration was approx. 5 times lower than that of SIgA, i.e. 20μg IgG/ml saliva and the inter-individual variation was lower than that observed for SIgA. The saliva-serum ratio for IgG isotypes IgG3/5 and IgG4/7 was also assessed in the sampled horses and this analysis showed that the saliva-serum ratio of IgG4/7 was in general approximately 4 times higher than that of IgG3/5. The large inter-individual variation in salivary SIgA levels observed for the normal healthy horses in the present study emphasises the need for a large number of observations when studying this parameter especially in a clinical setting. Moreover, our results also indicated that some of the salivary IgG does not originate from serum but may be produced locally. Thus, these results provide novel insight, and a base for further research, into salivary antibody responses of horses. |
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0165-2427 |
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Equine Behaviour @ team @ |
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6514 |
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Yarnell, K.; Hall, C.; Royle, C.; Walker, S.L. |
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Title |
Domesticated horses differ in their behavioural and physiological responses to isolated and group housing |
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Journal Article |
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Year |
2015 |
Publication |
Physiology & Behavior |
Abbreviated Journal |
Physiol.Behav. |
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143 |
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51-57 |
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Equine; Behaviour; Corticosterone; Housing |
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Abstract The predominant housing system used for domestic horses is individual stabling; however, housing that limits social interaction and requires the horse to live in semi-isolation has been reported to be a concern for equine welfare. The aim of the current study was to compare behavioural and physiological responses of domestic horses in different types of housing design that provided varying levels of social contact. Horses (n = 16) were divided equally into four groups and exposed to each of four housing treatments for a period of five days per treatment in a randomized block design. The four housing treatments used were single housed no physical contact (SHNC), single housed semi-contact (SHSC), paired housed full contact (PHFC) and group housed full contact (GHFC). During each housing treatment, adrenal activity was recorded using non-invasive faecal corticosterone metabolite analysis (fGC). Thermal images of the eye were captured and eye temperature was assessed as a non-invasive measure of the stress response. Behavioural analysis of time budget was carried out and an ease of handling score was assigned to each horse in each treatment using video footage. SHNC horses had significantly higher (p = 0.01) concentrations of fGC and were significantly (p = 0.003) more difficult to handle compared to the other housing types. GHFC horses, although not significantly different, had numerically lower concentrations of fGC and were more compliant to handling when compared to all other housing treatments. Eye temperature was significantly (p = 0.0001) lower in the group housed treatment when compared to all other treatments. These results indicate that based on physiological and behavioural measures incorporating social contact into the housing design of domestic horses could improve the standard of domestic equine welfare. |
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0031-9384 |
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Equine Behaviour @ team @ |
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5920 |
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Hartmann, E.; Søndergaard, E.; Keeling, L.J. |
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Title |
Identifying potential risk situations for humans when removing horses from groups |
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Journal Article |
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Year |
2012 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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136 |
Issue |
1 |
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37-43 |
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Equine; Group; Human; Handling; Injury |
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Removing a horse from its social group may be considered risky, both for the handler and the horse, because other horses can interfere in the catching process. The main aim of this study was to identify where and when these risk situations occur while removing a horse from its group. A potential risk situation was defined by the closeness of loose horses in the group or by any physical contact with them. Whether the number of horses following would be influenced by the social rank of the horse being led out, and whether more horses would follow to the gate when a larger proportion of the group was removed compared to when a single horse was taken out were also investigated. Thirty-two mares (1–2 years) were kept in groups of four. All horses were taken out of their home paddock twice alone (64 tests) and twice with a companion (32 tests). One handler (or two handlers when two horses were removed) was asked to approach (phase 1) and catch the target horse (phase 2), walk it to the centre of the paddock and remain stationary at a post for 30 s (phase 3), walk to the paddock entrance (phase 4) and through the gate (phase 5). The number of horses following, and the number of loose horses in proximity (<2 m, 2–5 m) to the target horse and handler was estimated, and horse–horse and horse–human interactions were recorded continuously for the five scoring phases. Significantly more loose horses were within 2 m of a single target horse during the phases approach (mean ± SD: 1.5 ± 0.8), catch (1.6 ± 0.9) and post (1.7 ± 0.7) than during walk (1.0 ± 0.5) and gate (1.1 ± 0.6). Rank did not influence the number of horses following to the gate (high rank: 2.4 ± 0.7; lower rank: 2.0 ± 1.0; P = 0.396) and interactions between horses were rare. A greater proportion of the loose horses followed when two horses (0.9 ± 0.2) were removed compared to when a single horse (0.7 ± 0.3) was taken out (P = 0.011). In conclusion, maintaining a distance to other horses in the group by reducing the time being relatively stationary, so giving loose horses fewer chances to approach, is likely to contribute to improved handler's safety. Removing a small proportion of the group may also decrease the probability of the other horses following. |
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0168-1591 |
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Equine Behaviour @ team @ |
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5491 |
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Uchiyama, H.; Ohtani, N.; Ohta, M. |
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Three-dimensional analysis of horse and human gaits in therapeutic riding |
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Journal Article |
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Year |
2011 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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135 |
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4 |
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271-276 |
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Equine-facilitated interventions; Horse gait; Human gait; Three-dimensional analysis; Horse riding |
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Therapeutic horse riding or hippotherapy is used as an intervention for treating individuals with mental and physical disabilities. Equine-assisted interventions are based on the hypothesis that the movement of the horse's pelvis during horseback riding resembles human ambulation, and thus provides motor and sensory inputs similar to those received during human walking. However, this hypothesis has not been investigated quantitatively and qualitatively. This study aimed to verify the hypothesis by conducting a three-dimensional analysis of the horse's movements while walking and human ambulation. Using four sets of equipments, we analysed the acceleration patterns of walking in 50 healthy humans and 11 horses. In addition, we analysed the exercise intensity by comparing the heart rate, breathing rate, and blood pressure of 127 healthy individuals before and after walking and horse riding. The acceleration data series of the stride phase of horse walking were compared with those of human walking, and the frequencies (in Hz) were analysed by Fast Fourier transform. The acceleration curves of human walking overlapped with those of horse walking, with the frequency band of human walking corresponding with that of horse walking. Exercise intensity, as measured by the heart rate and breathing rate, was not significantly different between horse riding and human walking. The levels of diastolic blood pressure were slightly higher during horse riding than during walking, but were lower during both conditions compared with those in normal conditions (P < 0.01). The present study shows that, although not completely matched, the accelerations of the horse and human walking are comparable quantitatively and qualitatively. Horse riding at a walking gait could generate motor and sensory inputs similar to those produced by human walking, and thus could provide optimum benefits to persons with ambulatory difficulties. |
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0168-1591 |
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Equine Behaviour @ team @ |
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5488 |
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Hockenhull, J.; Creighton, E. |
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Unwanted oral investigative behaviour in horses: A note on the relationship between mugging behaviour, hand-feeding titbits and clicker training |
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Journal Article |
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2010 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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127 |
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3-4 |
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104-107 |
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Equine; Horse; Titbits; Food rewards; Clicker training; Mugging behaviour |
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Unwanted oral investigative in horses has been anecdotally attributed to the practice of hand-feeding. Fears over such behaviour developing as a consequence of using food rewards, for example in clicker training, have been implicated as a common reason for not employing food-based positive reinforcement training techniques. This study used data generated as part of a larger research project, and explored associations between five common oral investigative behaviours and the practices of hand-feeding and clicker training. Data were from a convenience sample of UK leisure horse owners using two self-administered Internet surveys. Ninety-one percent of respondents reported giving their horse food by hand and this practice was significantly associated with three of the five oral investigative behaviours, licking hands (P = 0.006), gently searching clothing (P < 0.001) and roughly searching clothing (P = 0.003). Nipping hands and biting clothes were not associated with hand-feeding, suggesting that risk factors for these behaviours originate outside of this practice. Clicker training techniques were employed by 14% of respondents and their use was not associated with the incidence of any of the five oral investigative behaviours. These findings suggest that horse owners should not be deterred from using food-based positive reinforcement techniques with their horses, as fears that this practice will result in unwanted oral investigative behaviours from their horses appear unfounded. |
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0168-1591 |
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Equine Behaviour @ team @ |
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5183 |
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Burn, C.C.; Dennison, T.L.; Whay, H.R. |
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Relationships between behaviour and health in working horses, donkeys, and mules in developing countries |
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Journal Article |
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2010 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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126 |
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3-4 |
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109–118 |
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Animal welfare; Developing countries; Equine; Human-animal relationships; Inactivity; Sickness behaviour |
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Recent studies raise serious welfare concerns regarding the estimated 93.6 million horses, donkeys and mules in developing countries. Most equids are used for work in poor communities, and are commonly afflicted with wounds, poor body condition, respiratory diseases, parasites, dental problems, and lameness. Non-physical welfare problems, such as fear of humans, are also of concern. Interventions to improve working equine welfare aim to prioritise the conditions that cause the most severe impositions on the animals' subjectively experienced welfare, but data identifying which conditions these may be, are lacking. Here we describe a stage in the validation of behavioural welfare indicators that form part of a working equine welfare assessment protocol. Over 4 years, behavioural and physical data were collected from 5481 donkeys, 4504 horses, and 858 mules across nine developing countries. Behaviours included the animals' general alertness, and their responses to four human-interaction tests, using the unfamiliar observer as the human stimulus. Avoidance behaviours correlated significantly with each other across the human-interaction tests, with 21% of animals avoiding the observer, but they showed no associations with likely anthropogenic injuries. Over 13% of equids appeared [`]apathetic': lethargic rather than alert. Measures of unresponsiveness correlated with each other across the five tests, and were associated with poor body condition, abnormal mucous membrane colour, faecal soiling, eye abnormalities, more severe wounds, and older age, depending on the equine species. This suggests that working equids in poor physical health show an unresponsive behavioural profile, consistent with sickness behaviour, exhaustion, chronic pain, or depression-like states. |
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0168-1591 |
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Equine Behaviour @ team @ |
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5158 |
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Nagy, K.; Bodó, G.; Bárdos, G.; Bánszky, N.; Kabai, P. |
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Differences in temperament traits between crib-biting and control horses |
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Journal Article |
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2010 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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122 |
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1 |
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41-47 |
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Crib-biting; Stereotypic behaviour; Equine temperament; Trainability; Coping strategies |
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Recent studies have suggested that crib-biting in horses is associated with diminished capacity of learning or coping with stress. Such findings raise the question whether trainability, which is fundamentally important in practice, could also be affected by stereotypic behaviour. Trainability of a horse is difficult to assess in simple tests, however, it is reliably estimated by experienced riders. To assess trainability and other characteristics related to that, a questionnaire survey was conducted with the owners of 50 crib-biting and 50 control horses. Where possible, control horses were selected from the same establishment as crib-biters. Groups did not differ significantly regarding age, breed, gender, training level or usage. Principal component analysis revealed three main factors which can be labelled as [`]Anxiety', [`]Affability' and [`]Trainability'. The [`]Anxiety' factor consisted of the items [`]Nervousness', [`]Excitability', [`]Panic', [`]Inconsistent emotionality', [`]Vigilance', [`]Skittishness', and [`]Timidity'. [`]Affability' consisted of [`]Friendliness toward people', [`]Cooperation', [`]Docility' and [`]Friendliness toward horses'. [`]Trainability' involved [`]Concentration', [`]Trainability', [`]Memory', and [`]Perseverance'. Temperament traits were not affected by age, gender, breed or training level, but the usage of the horse and the presence of crib-biting behaviour had significant effects. Competition horses had lower level of [`]Anxiety' (p = 0.032) and higher level of [`]Trainability' (p = 0.068) than leisure horses. Crib-biting horses had significantly lower level of [`]Anxiety' than control horses (p < 0.001), while [`]Trainability' and [`]Affability' did not differ between groups (p = 0.823 and p = 0.543, respectively). Competition horses are more often exposed to novel environment and to frightening stimuli (e.g. colourful obstacles) than leisure horses and therefore might have also become more habituated to these types of stimuli. Coping with novel situation may be enhanced by defusing nervous behaviour by the more experienced riders of competition. Previous studies indicated crib-biting horses to be less reactive when challenged as compared to control horses. We suggest that the virtual calmness and lower nervousness of the crib-biting horses might be due to the passive coping style of these animals. [`]Affability' of horses might be more related to housing and management conditions than to crib-biting. Contrary to expectations, scores on [`]Trainability' had not coincided with the impaired learning of crib-biting horses reported in laboratory tests. However, previous behavioural tests on equine learning rarely had a direct relevance to the training abilities of the horses. Our results do not support crib-biting stereotypy to affect performance in training, which is a complex learning process involving cooperation and docility in the social environment. |
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Equine Behaviour @ team @ |
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5078 |
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Hartmann, E.; Christensen, J.W.; Keeling, L.J. |
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Title |
Social interactions of unfamiliar horses during paired encounters: Effect of pre-exposure on aggression level and so risk of injury |
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Journal Article |
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2009 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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Volume |
121 |
Issue |
3-4 |
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214-221 |
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Equine; Behaviour; Welfare; Mixing; Housing; Wounds |
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Group housing of horses is not widely applied in practice despite the welfare advantages of keeping animals socially rather than individually. In particular, concerns have been raised about the possible increased risk of injury and how to introduce a new horse into an established group. This study investigated two hypotheses: (1) pre-exposure of young horses in neighbouring boxes reduces the frequency of aggressive interactions when the same horses are subsequently put together in a paddock compared to horses without this previous box experience, (2) the occurrence of aggressive behaviour, in particular contact aggression in the paddock can be predicted after observing the horses' social interactions in neighbouring boxes. Danish Warmblood mares (n = 20), 2 years old, were kept in two groups of 10 horses. In total, 60 encounters were arranged whereby each horse was confronted pair-wise with six horses from the other group, three according to each treatment: treatment I--box (B) and subsequent paddock meeting (BP), and treatment II--only paddock meeting (P). Horses met in neighbouring boxes for 5 min and together in the same paddock for 10 min. The frequencies of aggressive and non-aggressive interactions were analysed from video recordings. Total aggression levels between BP and P did not differ, but [`]contact aggression', i.e. bite, kick, strike, push, tended to be lower in BP compared to P (median BP = 1, P = 2; p = 0.083) and there were less bites in BP than P (median BP = 0, P = 1; p = 0.050). Frequencies of [`]non-aggressive' interactions, e.g. friendly approach, nasal sniff, were lower in BP than P (median BP = 2.5, P = 10; p < 0.01). Results further revealed that [`]bite threat' performed in boxes correlated with [`]contact aggression' in the paddock (r = 0.46, p = 0.011). In conclusion, pre-exposure of young horses in neighbouring boxes may reduce [`]contact aggression', especially biting, in the paddock and [`]bite threat' shown in boxes may help to predict contact aggression when horses are later turned out together. The reduced non-aggressive interactions in the paddock in the BP test were probably a consequence of horses having exchanged these behaviours in the preceding B test. Exposing young horses in boxes next to each other may be a helpful tool before mixing them because horses meet in a safe environment that could assist in reducing the type of aggression where horses are most at risk of being injured. |
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Equine Behaviour @ team @ |
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5089 |
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Author |
Nagy, K.; Bodó, G.; Bárdos, G.; Harnos, A.; Kabai, P. |
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Title |
The effect of a feeding stress-test on the behaviour and heart rate variability of control and crib-biting horses (with or without inhibition) |
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Journal Article |
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2009 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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121 |
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2 |
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140-147 |
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Stereotypic behaviour; Crib-biting; Modified Forssell's operation; Cribbing collar; Equine welfare; Stress coping |
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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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0168-1591 |
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Equine Behaviour @ team @ |
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5091 |
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