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Burla,J.B.; Rufener, C.; Bachmann, I.; Gygax, L.; Patt, A.; Hillmann, E. |
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Effect of varying dimensions of the littered lying area on the lying behaviour of group-housed horses (Equus ferus caballus) |
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2015 |
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Proceedings of the 3. International Equine Science Meeting |
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Proc. 3. Int. Equine. Sci. Mtg |
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group housing, lying behaviour, littered lying area, rubber mats |
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Although horses can sleep while standing, recumbency is required for REM sleep and since all sleep stages must be completed for an entire sleep cycle, the opportunity for recumbency is essential for animal welfare. Observations on feral horses indicate a minimal lying duration of 30 min, preferably on a deformable and dry ground. In contrast to feral horses, lying behaviour in stabled horses is often affected by the dimensions of the provided lying area and rank.
In Switzerland, minimum requirements (MR) for the littered lying area are established by law to ensure animal welfare (BLV, 2008) (A/N: approximately match German recommendations (BMEL, 2009)). The aim of this study was to assess the adequacy of the dimensions of the minimum requirements for group-housed horses by investigating 38 horses in 8 groups. Further, hard rubber mats were provided supplementary in order to assess their suitability as an alternative to litter. Four treatments were each applied in randomised order:
– 0x MR: no litter + 1.5x MR with rubber mats
– 0.5x MR: 0.5x MR with litter + 1x MR with rubber mats
– 1x MR: 1x MR with litter; 0.5x MR with rubber mats
– 1.5x MR: 1.5x MR with litter + no rubber mats
For each treatment, after a habituation period of 8 days, lying behaviour was recorded (video, accelerometers) continuously for 72 hrs. Statistical analysis was performed using mixed effects models.
Regardless of the ground chosen, the duration of recumbency per 24 hrs was increasing with increasing dimensions of the littered area (F1,93 = 12.9, p = 0.0005; Fig. 1). Whereas the effect flattened from 1x to 1.5x MR, the duration spent on litter – a deformable ground – was increasing continuously (F1,62 = 23.1, p < 0.0001). Further, the proportion of lateral recumbency was increased with increased dimensions of the littered area (F1,79 = 12.3, p = 0.0007). Regarding the number of lying bouts, no differences were apparent between treatments providing litter, but recumbency occurred very seldom if only rubber mats were provided (F1,93 = 14.7, p = 0.0002). Further, low-ranking horses spent more lying bouts on rubber mats than high-ranking horses (F1,29 = 4.4, p = 0.04). Additionally, the larger the dimensions of the littered area the more horses were present in the lying area at the moment of lying down (F1,79 = 6.6, p = 0.01). Moreover, low-ranking horses showed considerably higher percentages of involuntarily terminated lying bouts than high-ranking horses if 0.5x and 1x MR were littered (F1,76 = 8.43, p = 0.005).
Although the indicated minimal lying duration was averagely performed, large individual differences occurred and at least 8% were lying down less than 30 min per 24 hrs in every treatment. Further, the inclusion of social parameters indicated a beneficial effect of an exceedance of the minimum requirements especially for low-ranking horses. Therefore, the minimum requirements established by Swiss law can be stated as adequate but should be perceived as minimum and not optimum dimensions. |
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Burla,J.B. |
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Equine Behaviour @ team @ |
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5879 |
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Malavasi, R.; Huber, L. |
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Referential communication in the domestic horse (Equus caballus): first exploration in an ungulate species |
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Conference Article |
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2015 |
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Proceedings of the 3. International Equine Science Meeting |
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Proc. 3. Int. Equine. Sci. Mtg |
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domestic horse, referential communication, human-horse communication, intentionality |
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An important question in the study of animal communication is whether non-human animals are able to produce communicative gestures, i.e. nonvocal bodily actions directed to a recipient, physically ineffective but with a meaning shared in the social group [1]. Passive gestures are instrumental, tuned to the mere presence/absence of others, whereas active informers recognize receivers as communicative agents and activate shared-attention mechanisms for identifying their attentional state (SAM [2]; e.g. Schwab and Huber [3]). Six operational criteria must be evaluated to classify a signal as referential and intentional [4]: (1) alternative gazes between the partner and the target; (2) apparent attention-getting behaviours are deployed; (3) an audience is required to exhibit the behaviour; (4) the attentional status of an observer influences the propensity to exhibit behaviours; (5) communication is persistent and (6) there is elaboration of communicative behaviour when apparent attempts to manipulate the partner fail. Dogs [5] and non-human primates (reviewed in Liebal and Call [6]) can tune a human receiver’s attention to the object of interest by combining directional and attention-getting signals, such as turning the head or body, gazing to the receiver, and/or establishing eye contact. Research on other species is scarce.
Horses rely on humans to survive in domestic settings and may have evolved skills for communicating flexibly with them [7]. Horses understand human attentional cues (such as body and head orientation, eyes opened/closed) [8], permanent pointing [9] and, to some extent, gazing [10]. Here we tested the ability of 14 outdoor, herd-living domestic horses to communicate referentially with a human partner about the location of a desired target, a bucket of food out of reach. After the baiting of two buckets placed in opposite, unreachable locations were shown by the experimenter, the subject would walk to one of the two buckets. Because approaching a bucket would reveal that the food is out of reach, we expected the horse to look back to the experimenter, then to the bucket, and alternate this gazing several times to indicate its intention. To test whether our prediction is correct and alternate gazing is indeed the result of the horse's referential communication, we video-recorded the behaviour of the subjects in the test (FORWARD) and three control conditions: (1) FORWARD: experimenter oriented to the center of the arena, (2) BACK: experimenter backward oriented in respect to the arena, (3) ALONE: experimenter absent, (4) MANY: as FORWARD plus a familiar human oriented to the subject behind the bucket (Figure 1). We used a conservative criterion of back gazing by considering only turning the head back more than 90 degrees. The results confirmed our prediction. The horses alternated gazes between the partner and the buck significantly more often in the FORWARD than in all the other conditions (Table 1), thus satisfying operational criteria #1, #3 and #4. They also alternated head nods with gazes to the partner significantly more often during the FORWARD condition. We thus considered head nods not an instrumental signal of arousal, but an attention-getting behaviour with communicative function. Subjects used both head nods and neck stretched toward the buck more often in the FORWARD than in the BACK and the ALONE conditions, thus satisfying criteria #2, #3 and #4. In condition MANY, the frequency of head nods did not differ from condition FORWARD, probably because nods were directed to the additional partner behind the buck. This also satisfies criteria #4. The horses gazed to the partner most often in the FORWARD than in the BACK and the MANY conditions, but not in the ALONE. In this condition, subjects could observe the partner walking further from the test arena. To test for the different functions of gazes in presence and in absence of the partner, we compared their average duration between the two conditions: the significantly longer duration of gazes when the subject was alone suggests the instrumental monitoring function of gazes in this experimental condition.
Altogether, the findings suggest that domestic horses possess the ability to use referential communication in an interspecific context, but additional analyses are needed to test for operational criteria #5 and #6. Flexible and voluntary use of communicative signals reveal sophisticated cognitive processes involved in the strategic emission of these signals, and the finding of referential communication skills in an ungulate species forces us to reconsider the evolutionary path of intelligence. Furthermore, ungulates are used intensively by humans (transportation, meat, agriculture, leisure activities), and their welfare is often compromised. Determining whether ungulates can communicate their needs and preferences is paramount to a proper ethical management. |
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Malavasi, R. |
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Equine Behaviour @ team @ |
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5876 |
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Author |
Rubenstein, D. |
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Title |
Social Networks: Linking Form with Function in Equid Societies |
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Conference Article |
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2015 |
Publication |
Proceedings of the 3. International Equine Science Meeting |
Abbreviated Journal |
Proc. 3. Int. Equine. Sci. Mtg |
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Animal societies develop from interactions and relationships that occur among individuals within populations. The fundamental tenet of behavioral ecology is that ecological factors shape behavior and determine the distribution and associations of individuals on landscapes. As a result, different social systems emerge in different habitats and under different environmental conditions. Since characterizing social systems depends on time and motion studies of individual actions and interactions that are often bilateral, such characterizations are often coarse-grained. If social relationships can be characterized using social networks, however, seemingly similar social organizations often reveal informative differences in terms of deep structure. Thus social network theory should be able to provide insights in to the connections between social form and function. This talk will explore how the network structures of horses, zebras and asses can provide novel insights into the functioning of animal societies with respect to the spread of memes, genes and diseases. |
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Rubenstein, D. |
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Xenophon Publishing |
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Wald |
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Krueger, K. |
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Proc. 3. Int. Equine. Sci. Mtg |
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in prep |
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978-3-95625-000-2 |
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Equine Behaviour @ team @ |
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5797 |
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Ahmadinejad , S.M.; Pishkar, J.; Bahmen, M. |
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Comparisons of behavioral and physiological state in Caspian pony before and after stress |
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2015 |
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Proceedings of the 3. International Equine Science Meeting |
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Proc. 3. Int. Equine. Sci. Mtg |
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Behavioral scores (BS) offer an non-invasive, objective and easy to use way of assessing welfare in horses. Their development has, however, largely focused on behavioral reactions to stressful events (often induced), and so far no use of physiological measures has been made to underpin and validate the behavioral measures in the Caspian ponies. This study aimed to develop a physiologically validated scale of behavioral indicators of stress for the purpose of welfare logically validated scale of behavioral and physiological data assessment in the stabled Caspian ponies. To achieve this , behavioral and physiological data were collected from 16 Caspian ponies that underwent routine husbandry procedures. The ponies were divided into two groups, a control and a treatment group (8 each). The ponies in the treatment group took part in a 700 meter race. Analysis of the behavioral data were undertaken by a panel of equestrian industry professionals. Physiological measures (salivary and serum cortisol level) were significantly correlated with the behavioral scores confirming that the scores were meaningful and reflected the physiological stress. The scores offer an easy to use tool for rapid, reliable non-invasive welfare assessment in Caspian ponies, and reduce the need for potentially invasive physiological measures. |
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Ahmadinejad , S.M. |
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Xenophon Publishing |
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Wald |
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Krueger. K. |
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978-3-95625-000-2 |
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Equine Behaviour @ team @ |
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5891 |
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Hollenhorst, H.; Weil, S.; Krueger, K. |
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Title |
Innovative behavour in horses (Equus caballus) |
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Conference Article |
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2015 |
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Proceedings of the 3. International Equine Science Meeting |
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Proc. 3. Int. Equine. Sci. Mtg |
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innovative behavior, age, housing system, horse (Equus caballus) |
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Contrary to the widely-spread assumption that horses just have restricted cognitive capacities and are not very flexible in their behaviors, we showed that horses display innovative behavior and even make use of tools (Krueger 2015, Krueger et al. 2015). These findings derive from a database (http://innovative-behaviour.org/) the Equine behavior team managed in the past two years. Some horses did not only show single innovations, but several different innovations. The number of innovations per individual varied from 1 to 10. 20 % of all innovative horses in the database showed more than one innovation. These individuals can be called the ‘true innovators’. Moreover innovations were dependent on age. Young horses were more innovative than older ones, whereby horses at the age of five to nine years were the most innovative. When considering the housing system innovative horses in a single housing (inside box, outside box, paddock box) had a slight majority towards horses in group housing (open stable, active stable, pasture day and night). But given the fact that ratings on housing system frequencies state 95% of the horses to be kept in individual housing, innovations in individual housing are rare. Nevertheless, horses kept in inside boxes without a window, opened doors more often than all other horses. Aside from this effect, housing systems did not trigger the frequency of innovative behavior. Innovations for gaining freedom and innovations in general were widespread among horses with daily access to pasture and daily contact with conspecifics. Innovations for gaining food were not more likely to occur in horses that were fed little amounts of roughage. In conclusion, the housing of horses does not seem to be the primary catalyst for developing innovative behavior in horses. What makes a “true innovator” in horses, in addition to age, remains to be seen. |
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Hollenhorst, H. |
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Xenophon Publishing |
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Wald |
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; Krueger, K. |
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Proc. 3. Int. Equine. Sci. Mtg |
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978-3-95625-000-2 |
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Equine Behaviour @ team @ |
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5896 |
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Briefer Freymond, S.; Piovesana, L.; Briefer. E. F.; Beuret. S.; Zuberbühler, K.; Bshary, R.; Bachmann, I. |
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Crib-biting behaviour of horses: stress and learning capacity |
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Conference Article |
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2015 |
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Proceedings of the 3. International Equine Science Meeting |
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Proc. 3. Int. Equine. Sci. Mtg |
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Crib-biting is a stereotypy in horses that is potentially linked to both chronic stress and genetic predisposition. Chronic stress can cause neurobiological changes such as alteration of the dopaminergic modulation of the basal ganglia [1]. These neurobiological changes could alter and modify the learning profile of the horses [2,3]. We tested 19 crib-biters and 18 non-crib-biting horses (controls) in five challenging spatial tasks, in order to test if differences in dopaminergic modulation impair learning capacities. The tests were performed in two time periods, in a small arena (8 x 10 m) that was familiar to the horses. For each trials, the horses were led to the start zone in front of a four-meter-long solid fence and were then left alone in the arena. Their task was then to find a bucket containing food, which was situated in different positions around the fence, depending on the tests. The time to reach the food bucket, the trajectory taken by the horse (left or right side of the fence) and the ECG trace were recorded continuously. Additionally, salivary cortisol was collected before the tests (baseline), after the first time period, and after the second time period. We found that crib-biters and controls behaved similarly during the learning tasks. However crib-biters that did crib-bite on the solid fence during the task (group A; 10 horses) behaved differently than crib-biters that did not crib-bite (group B; 9 horses) and controls (group C; 18 horses) for some tests, in their trajectory or time to reach the bucket. These differences are more likely explained by the time taken to crib-bite, than by differences in learning capacity. We did not find any difference between groups in their heart-rate variability (RMSSD). Yet, we found a difference in salivary cortisol after the first time period between groups A, B and C. Indeed, the crib-biters that did not crib-bite had higher salivary cortisol values than all the other horses (mean±SE: A, 0.51±0.16ng/ml, B, 0.78±0.17ng/ml, C, 0.59±0.20ng/ml; Linear mixed model (LMM), p<0.05). Our results suggest that crib-biting horses that did not crib-bite during the learning tasks were more stressed than all other horses. This difference could be due to higher stress sensitivity in crib-biters, which could be reduced by the opportunity to crib-bite. These results replicate our previous findings testing differences in cortisol levels between crib-biters and control horses during an ACTH challenge test. Therefore, crib-biting behaviour might be a coping strategy helping stereotypic horses to reduce their stress during frustrating situations [4].
Keyword:
stereotypy, chronic stress, learning task |
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stereotypy, chronic stress, learning task |
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Briefer Freymond, S. |
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Equine Behaviour @ team @ |
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5878 |
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Medill, S.A; Janz, D.M.; McLoughlin, P.D. |
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Hair testosterone and cortisol concentrations and their relationships to physiological and social status in feral horses (Equus ferus caballus) |
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Conference Article |
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2015 |
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Proceedings of the 3. International Equine Science Meeting |
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Proc. 3. Int. Equine. Sci. Mtg |
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Determining steroid hormone concentrations in hair has been frequently performed in humans, and increasingly in wildlife and domestic animals. Hair hormone concentrations may provide insight on how individuals are responding to their physiological condition or social situation. Cortisol is most frequently measured in hair as a biomarker of long-term stress, while testosterone may be linked with reproductive status in males. These hormones are commonly measured in substances that reflect either current (e.g. blood) or very recent (e.g. saliva, urine, feces) circulating levels. However, these hormones are also incorporated into hair during hair growth and provide a chronological record of circulating hormone levels. Thus, analysis of steroid hormones in hair provides a much longer representation of an animal’s endocrine status than other tissues frequently targeted for non-invasive monitoring.
The feral horse (Equus ferus caballus) population on Sable Island, Nova Scotia, Canada, has been annually censused during the mid-late summer since 2008 to track individual life histories and population dynamics. We collected tail hair (n = 144 females, n = 162 males) from known individuals either opportunistically or from natural or artificial snags to investigate how hair cortisol and testosterone might be associated with physiological state (e.g. lactating vs. non-lactating, body condition, age), as well as their social situation (e.g. dominant band stallion, subordinate band stallion, or bachelor) and measures of sociality. The proximal 5 cm of hair (excluding first 4mm or root region) were ground to a fine powder and hormones extracted with methanol and analyzed by using enzyme-linked immunoassay.
Preliminary analyses of the data showed a general sex based difference in hair cortisol concentrations (females lower than males; t = 3.16, df = 317, P=0.002). Among females, the presence of nursing foals was accompanied by an increase in hair cortisol (z = 2.93, df =140, P = 0.003); however, no significant difference was found in hair cortisol concentrations based on sex of the foal (t = -0.06, df = 82, P = 0.95). Horses in poor body condition tended to have higher hair cortisol than those in good or excellent condition (slope= -0.203, df = 312, P = 0.003). We also observed an increased concentration of hair cortisol as horses increased in age from 3-6 or entered into reproductive maturity. Adult male dominant band stallions did not have significantly less cortisol than bachelors or subordinate stallions but these three groups were significantly greater than young males (aged 3 and 4) who generally do not challenge the older males for reproductive opportunities.
Additionally, we looked at hair testosterone concentrations for n=46 males. Testosterone is known to influence traits and behaviours that enhance sexual selection. Often there is an inverse relationship between cortisol levels and testosterone; in particular, being able to maintain high testosterone and not have elevated cortisol related to the metabolic costs of sexual trait production ensures that traits or behaviours honestly signal the quality of the individual. For this reason we’d expect to see band stallions (those males in a position to mate) have a lower value in the ratio Cortisol: Testosterone. Early indications suggest we see this phenomenon in feral horses.
As a relatively new approach in wildlife research, the use of hair hormone analysis shows promise in contributing to our understanding of physiological aspects of sexual selection and other processes. Additionally, hair hormone analysis may have applications in advancing knowledge of animal husbandry and in particular, welfare. |
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Medill, S.A. |
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Equine Behaviour @ team @ |
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5870 |
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Ahmadinejad , S.M.; Pishkar, J.; Anisi, T.; Babayee, B. |
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Verbal expressions of the horses to the pain |
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Conference Article |
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2015 |
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Proceedings of the 3. International Equine Science Meeting |
Abbreviated Journal |
Proc. 3. Int. Equine. Sci. Mtg |
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For communicating with the environment (other horses, foals, owner, etc.), horses have to use different methods. In contrast with the human, for whom talking is the most important way of communication, the horses can’t talk. In the years 1990s and before, the imagination was such that the infants do not express verbally to the pain either (like horses).
To communicate, horses use their body language. Vocalization (if not the body language), seems to be the most important way of communication in horses, though it seems they use the same tone when exposing to different events.
In this study which was performed in collaboration with the electronic institute of Sharif technical university (the top most technical university of the country), totally 25 horses were used. The horses were exposed to different events (hunger, pain, loneliness, mating, parturition and separation of the 1-2 weeks old foals from their dams). The verbal expressions of the horses were studied using spectrogram.
The results of this study showed that there were significant differences between the spectrograph of the voices of the horses, exposed to hunger, and the separation of the foals from dams. This was the same (no significant differences) when horses were exposed to loneliness, separation and hunger. There were no significant differences between the verbal expressions of the horses while exposed to mating, parturition.
One of the most important reasons why the horses do not have verbal expressions when exposing to the pain, might be the absence of the part of the brain, responsible for the pain interpretation. Morse research has to be performed to prove this. |
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Ahmadinejad , S.M. |
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Xenophon Publishing |
Place of Publication |
Wald |
Editor |
Krueger. K. |
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978-3-95625-000-2 |
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Equine Behaviour @ team @ |
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5890 |
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Author |
Jacquot, M.; Grosjean, A., Emrot, C.; Van-Erck-Westergem, E.; Schwartz, C.; Tomberg, C. |
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Effects of a walk phase at the warm up onset on physiological and behavioural parameters of ridden horses (Equus caballus) |
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2015 |
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Proceedings of the 3. International Equine Science Meeting |
Abbreviated Journal |
Proc. 3. Int. Equine. Sci. Mtg |
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For human athletes, any physical performance classically begins with a warm-up including a phase of cardio-respiratory activation as well as a phase of neuro-muscular mobilization. For the equine athletes similar routines are widely followed despite limited scientific studies addressing its effectiveness. Although physiological bases are of a similar nature in humans and horses, the latter are showing a different “telos” related to their status of prey. Indeed, their survival depends on their ability to detect any predator or potential danger in their environment and to estimate the need to run away. Therefore warm-up techniques should be adapted to this specificity and we added a walk phase preceding the usual warm-up during which the horse was allowed to visually assess its environment without constraint imposed by its rider, reins being long. We assumed that this walk phase would allow a more relaxed mental state and then a decrease in cognitive resources involved in environmental monitoring. Consequently this would release cognitive resources then available for the communication between the horse and the rider.
The autonomous nervous system (ANS) regulates the heart rate via the sympathetic and the parasympathetic nervous systems. The sympathovagal balance and the heart rate variability are considered as good indicators of an acute stress state as well as emotional states. These measures are frequently used to estimate well-being in animals. During the walk phase, we observed a significant decrease of the heart rate (HR), a significant increase of its variability and a significant modification of the sympathovagal balance in favour of a higher contribution of the parasympathetic control. We also measured the non-linear correlation dimension which reflects the degree of freedom of a system. To the best of our knowledge, this method was used until now only in humans to whom it was shown that its decrease is related to stressful events and is associated with a bad prognosis for survival. During the walk phase in horses, the dimension of correlation of cardiac activity was increased.
From a behavioural point of view, we observed a significant decrease of side movements related to head orientation as well as significant changes in ears position, the latter being preferentially directed forwards at the beginning of the walk phase and laterally at the end. As previous research suggests that ears position and head orientation can be involved in attentional mechanisms in horses, our results suggest a decrease of the attention focused on the environment. Furthermore, a significant lowering of the neck was observed as well as a significant decrease of behaviours related to stress.
To conclude our results suggest that a walk phase preceding the usual warm-up of horses would contribute to improve their mental state in favour of more relaxation and of a decrease of the attentional resources invested in environmental monitoring. |
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Corporate Author |
Jacquot, M. |
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Xenophon Publishing |
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Wald |
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; Krueger, K. |
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978-3-95625-000-2 |
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Equine Behaviour @ team @ |
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5897 |
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Author |
Bouskila, A.,; Lourie,E.; de Vries, H.; Hermans, Z.M .; van Dierendonck, M. |
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Sex, but not relatedness nor age, affect the social network of horses in a semi-natural reserve |
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Conference Article |
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2015 |
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Proceedings of the 3. International Equine Science Meeting |
Abbreviated Journal |
Proc. 3. Int. Equine. Sci. Mtg |
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For some time now, social network analysis provides tools to describe concisely the social structure of animals. Nevertheless, the factors that shape the social network and determine the frequency of different types of affiliations are often unclear. For example, the effects of relatedness on social networks have been only studied on relatively few species. Both among social or sub-social species the effects ranged from no effect to a substantial effect. Here we test the hypothesis that relatedness has an effect on the social network of horses (Equus caballus) that live freely in semi-natural conditions, and specifically, that individuals with high values of relatedness to their neighbors in the network will have fewer links Wolf et al, 2011). This hypothesis is based on the notion that related individuals have common interests and are likely to assist each other, and may need less links to other, unrelated individuals. In addition, individuals of similar age and sex are likely to have common needs, and thus are more likely to be associated. We thus tested a second hypothesis: homophily to individuals with similar age and sex will have a role in determining the associations within the social network. The field study was conducted on 27 horses in the Blauwe Kamer reserve in the Netherlands (1.1 sq km). We videotaped horses and their groups and used the information from 22 horses, after excluding the five foals from the analysis. Relatedness was calculated from the pedigree, which was based on parentage, determined by DNA analysis. The social network was constructed based on spatial proximity data. We assessed the influence of relatedness, age-homophily and sex-homophily on the network structure with Multiple Regression Quadratic Assignment Procedure (MRQAP) (Krackhardt 1988), with the R package sna. The results show that there was no significant effect of relatedness on the network, nor an effect of age-homophily. Nevertheless, we found a significant effect of sex-homophily, the tendency of individuals to associate with individuals of the same sex. We argue that the lack of a relatedness effect is not likely to have been caused due to the inability to detect who is kin. The structured social system in horses includes strong associations between often unrelated individuals, alongside with young individuals leaving their natal harem as part of the inherent inbreeding avoidance (Linklater & Cameron 2009; Boyd et al. in press). The significant effect of sex-homophily could stem from the protection females get from associations with other females, e.g., in the reduction of harassment from males. Previous studies on feral horses showed that mares that were better connected with other females in their harem benefited from higher survival rates to their foals (Cameron et al. 2009). The associations among bachelor males could also contribute to the strength of the sex-homophily effect.
In order to generalize from our results, one needs to examine additional populations of horses, because the conditions in the Blauwe Kamer reserve may not be representative, mainly due to the limited opportunity for dispersal in a restricted area.
Keywords: Long-term affiliation; spatial proximity, kin detection
Boyd, L., Scorolli, A. L., Nowzari, H., & Bouskila, A. (2016). Chapter 2: Social organization. In J. I. Ransom, & P. Kaczensky (Eds.), Wild equids. Baltimore, Maryland (in press): The Johns Hopkins University Press.
Cameron, E. Z., Setsaas, T. H., & Linklater, W. L. (2009). Social bonds between unrelated females increase reproductive success in feral horses. Proceedings of the National Academy of Sciences, 106(33), 13850-13853
Krackhardt, D. (1988). Predicting with networks: Nonparametric multiple regression analysis of dyadic data. Social Networks, 10(4), 359-381.
Linklater, W. L., & Cameron, E. Z. (2009). Social dispersal but with philopatry reveals incest avoidance in a polygynous ungulate. Animal Behaviour, 77(5), 1085-1093.
Wolf, J. B., Traulsen, A., & James, R. (2011). Exploring the link between genetic relatedness r and social contact structure k in animal social networks. The American Naturalist, 177(1), 135-142. |
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Bouskila, A. |
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Equine Behaviour @ team @ |
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5881 |
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