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Krueger, K., Schneider, G., Flauger, B., & Heinze, J. (2015). Context-dependent third-party intervention in agonistic encounters of male Przewalski horses. Behav. Process., 121, 54–62.
Abstract: Abstract One mechanism to resolve conflict among group members is third party intervention, for which several functions, such as kin protection, alliance formation, and the promotion of group cohesion have been proposed. Still, empirical research on the function of intervention behaviour is rare. We studied 40 cases of intervention behaviour in a field study on 13 semi-wild bachelor horses (Equus ferus przewalskii) in (a) standard social situations, and (b) when new horses joined the group (i.e. introductions). Only interventions in agonistic encounters were analysed. Eight of 13 animals directed intervention behaviour toward threatening animal in agonistic encounters of group members. One stallion was particularly active. The stallions did not intervene to support former group mates or kin and interventions were not reciprocated. In introduction situations and in standard social situations, the interveners supported animals which were lower in rank, but targeted, threatening animals of comparable social rank. After introductions, stallions received more affiliative behaviour from animals they supported and thus appeared to intervene for alliance formation. In standard social situations, interveners did not receive more affiliative behaviour from animals they supported and may primarily have intervened to promote group cohesion and to reduce social disruption within the group.
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Huebner, F., & Fichtel, C. (2015). Innovation and behavioral flexibility in wild redfronted lemurs (Eulemur rufifrons). Anim.Cogn., 18(3), 777–787.
Abstract: Innovations and problem-solving abilities can provide animals with important ecological advantages as they allow individuals to deal with novel social and ecological challenges. Innovation is a solution to a novel problem or a novel solution to an old problem, with the latter being especially difficult. Finding a new solution to an old problem requires individuals to inhibit previously applied solutions to invent new strategies and to behave flexibly. We examined the role of experience on cognitive flexibility to innovate and to find new problem-solving solutions with an artificial feeding task in wild redfronted lemurs (Eulemur rufifrons). Four groups of lemurs were tested with feeding boxes, each offering three different techniques to extract food, with only one technique being available at a time. After the subjects learned a technique, this solution was no longer successful and subjects had to invent a new technique. For the first transition between task 1 and 2, subjects had to rely on their experience of the previous technique to solve task 2. For the second transition, subjects had to inhibit the previously learned technique to learn the new task 3. Tasks 1 and 2 were solved by most subjects, whereas task 3 was solved by only a few subjects. In this task, besides behavioral flexibility, especially persistence, i.e., constant trying, was important for individual success during innovation. Thus, wild strepsirrhine primates are able to innovate flexibly, suggesting a general ecological relevance of behavioral flexibility and persistence during innovation and problem solving across all primates.
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Aurich, J., Wulf, M., Ille, N., Erber, R., von Lewinski, M., Palme, R., et al. (2015). Effects of season, age, sex and housing on salivary cortisol concentrations in horses. Domest. Anim. Endocrinol., .
Abstract: Abstract Analysis of salivary cortisol is increasingly used to assess stress responses in horses. Since spontaneous or experimentally induced increases in cortisol concentrations are often relatively small for stress studies proper controls are needed. This requires an understanding of factors affecting salivary cortisol over longer times. In this study, we have analysed salivary cortisol concentration over 6 mo in horses (n = 94) differing in age, sex, reproductive state and housing. Salivary cortisol followed a diurnal rhythm with highest concentrations in the morning and a decrease throughout the day (P < 0.001). This rhythm was disrupted in individual groups on individual days; however, alterations remained within the range of diurnal changes. Comparison between months showed highest cortisol concentrations in December (P < 0.001). Cortisol concentrations increased in breeding stallions during the breeding season (P < 0.001). No differences in salivary cortisol concentrations between non-pregnant mares with and without a corpus luteum existed. In stallions, mean daily salivary cortisol and plasma testosterone concentration were weakly correlated (r = 0.251, P < 0.01). No differences in salivary cortisol between female and male young horses and no consistent differences between horses of different age existed. Group housing and individual stabling did not affect salivary cortisol. In conclusion, salivary cortisol concentrations in horses follow a diurnal rhythm and are increased in active breeding sires. Time of the day and reproductive state of the horses are thus important for experiments that include analysis of cortisol in saliva.
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Giles, S. L., Nicol, C. J., Harris, P. A., & Rands, S. A. (2015). Dominance rank is associated with body condition in outdoor-living domestic horses (Equus caballus). Applied Animal Behaviour Science, .
Abstract: Abstract The aim of our study was to explore the association between dominance rank and body condition in outdoor group-living domestic horses, Equus caballus. Social interactions were recorded using a video camera during a feeding test, applied to 203 horses in 42 herds. Dominance rank was assigned to 194 individuals. The outcome variable body condition score (BCS) was recorded using a 9-point scale. The variables age and height were recorded and considered as potential confounders or effect modifiers. Results were analysed using multivariable linear and logistic regression techniques, controlling for herd group as a random effect. More dominant (p = 0.001) individuals generally had a higher body condition score (p = 0.001) and this association was entirely independent of age and height. In addition, a greater proportion of dominant individuals fell into the obese category (BCS >= 7/9, p = 0.005). There were more displacement encounters and a greater level of interactivity in herds that had less variation in age and height, lending strength to the hypothesis that phenotypic variation may aid cohesion in group-living species. In addition there was a strong quadratic relationship between age and dominance rank (p < 0.001), where middle-aged individuals were most likely to be dominant. These results are the first to link behavioural predictors to body condition and obesity status in horses and should prompt the future consideration of behavioural and social factors when evaluating clinical disease risk in group-living animals.
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Wyss, C. (2015). Does housing in a „social box“ change faecal cortisol metabolites concentration in stallions? In Proceedings of the 3. International Equine Science Meeting.
Abstract: In order to improve the housing conditions of stallions in individual boxes by offering a possibility to have more social contact, the Swiss national stud farm tested a new box system for horses, allowing increased physical contact with the neighbouring stallion. The aim of this part of the study was to investigate whether this type of housing system (named “social box”) potentially induces a change in stress reactions in stallions compared to conventional boxes. Therefore faecal cortisol metabolite (FCM) concentration was measured as a non-invasive parameter to assess endocrine responses related to this new environment.
Four groups each consisting of eight adult Freiberger breeding stallions were included in the test design. Every stallion spent three weeks in a conventional box and in a social box respectively (cross-over design). The conventional box consisted of a separation wall with a lower opaque part and an upper part with vertical barriers (5 cm between barriers), allowing visual and olfactory contact but strongly limiting tactile contact. The separating wall of the social box consisted of two lateral sections, one part being opaque to the ceiling and the second part consisting of vertical barriers (30 cm between barriers), allowing the horse to have physical contact with its neighbour or to avoid it.
In horses, FCM concentration reflects an average level of circulating cortisol over a period of approximatively 24h. Faecal samples were collected the day following integration in social / conventional boxes, reflecting the potential stress induced by increased social interactions during the integration. In order to asses potential chronical stress, faeces samples were also collected in week one, two and three after the integration into the social / conventional box (in total: 4 samples per horse and housing system). The samples were immediately stored at -20°C until they were analysed. The samples were not analysed in the laboratory until the end of the experiment, therefore the duration of conservation in the freezer varied from 40 to 429 days.
A considerable percentage of data from groups 1 and 2 was below the detection limit (<0.8 ng/g) (Tab. 1). Thus the statistical analysis was conducted with the FCM concentration from groups 3 and 4 (n horses = 16) which contained no values below the detection limit.
Tab. 1: Details about FCM values and storage time for the 4 groups of stallions
Group Storage duration [d] Proportion of data below the detection limit (<0.8 ng/g) Mean [ng/g] Median [ng/g]
Group 1 384-429 55.6 % 2.2 0
Group 2 315-360 25.5 % 5.8 6.3
Group 3 41-79 0.0 % 8.7 8.0
Group 4 40-85 0.0 % 5.8 5.4
Despite the impressive social interactions observed between the stallions directly after being introduced into the social boxes, we did not find any differences in FCM concentration between the stallions being introduced into the conventional box and the social box on the day of integration (social box: n samples = 16, mean±SD: 6.9±4.7 ng/g; conventional box: n samples = 16, mean±SD: 9.0±11.2 ng/g; Wilcoxon signed rank test V = 70, p = 0.94).
Overall the samples taken during integration and in week one, two and three did not show evidence of changes in FCM concentration in either housing system over a longer period of time (social box: n samples = 64, mean±SD: 7.9±6.2 ng/g; conventional box: n samples = 64, mean±SD: 6.6±3.4 ng/g; Linear mixed model (LMM), p = 0.56).
Our results suggest that the possibility of having physical contact with a conspecific does not induce changes in FCM concentration in breeding stallions. The considerable percentage of values below the detection limit in groups 1 and 2 seemed to correlate with the increasing duration of storage before analysis. During the IESM Network Meeting 2015, we would like to discuss possible methodological issues and the possibilities to correctly integrate these low values in the statistical analysis.
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Medill, S. A., Janz, D. M., & McLoughlin, P. D. (2015). Hair testosterone and cortisol concentrations and their relationships to physiological and social status in feral horses (Equus ferus caballus). In Proceedings of the 3. International Equine Science Meeting.
Abstract: 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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Fuchs, C., Kiefner, C., Erhard, M., & Wöhr, A. C. (2015). Narcolepsy – or REM-deficient? In Proceedings of the 3. International Equine Science Meeting.
Abstract: Narcolepsy is a neurological sleep disorder characterized by excessive daytime sleepiness, cataplexy (loss of muscle tone), sleep paralysis and hypnagogic hallucinations, also called the „tetrad of narcolepsy“. Although the pathogenesis is not completely understood, the disorder is well described in humans and it has been shown that a lack of the hormone hypocretin (orexin) synthesized in the hypothalamus is crucial. Narcolepsy with cataplectic attacks has also been reported in dogs, horses, cattle (STRAIN et al., 1984) and a lamb (WHITE und DE LAHUNTA, 2001).
In dogs up to 17 breeds have been shown to be affected sporadically, while familial forms occur in dobermans, labrador retrievers and dachshounds (TONOKURA et al., 2007). In horses there appear to be two syndroms (HINES, 2005), the first in which animals are affected within a few days after birth (possibly a familial form, reported in Suffolk, Shetland ponies, Fell ponies, Warmbloods, Miniature Horse foals (MAYHEW, 2011), Lipizzaner (LUDVIKOVA et al., 2012) and Icelandic horses (BATHEN‐NÖTHEN et al., 2009)) and the second in which animals are affected as adults (adult-onset narcolepsy).
It has been shown that both forms of canine narcolepsy are associated with a deficit in hypocretin/orexin neurotransmission (LIN et al., 1999). In the horse a similar etiology is suspected, but so far there are no studies to support this hypothesis.
The cataplectic attacks in humans and dogs occur during excitement or emotional stimulation such as laughing in humans or eating and playing in dogs. In contrast, the cataplectic or sleep attacks in adult horses happen almost exclusively while resting. The collapses observed in equines vary from drowsiness with hanging of the head, swaying, buckling at the knees or total collapse (see fig.1). Affected horses often show injuries and scars at the dorsal fetlocks, dorsal knees or at the face and the lips. ALEMAN et al. (2008) describe some of the suspected adult-onset narcolepsy cases as possible examples of sporadic idiopathic hypersomnia instead of true narcolepsy.
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König von Borstel, U., Kienapfel, K., McLean, A., Wilkins, C., Evans, D., & McGreevy, P. (2015). Hyperflexing the horse‘s neck: a cost-benefit and meta-analysis. In Proceedings of the 3. International Equine Science Meeting.
Abstract: In ethical discussions, a cost-benefit analysis requires that welfare costs associated with an activity can be reliably estimated and balanced against the potential benefits of the activity to both humans and animals. The current study applies a meta-analysis to the peer-reviewed evidence for costs and benefits of hyperflexion of the neck in horses; a practice that has attracted enormous public and scientific scrutiny over the past 15 years. A literature review identified 55 studies dealing with horses’ head and neck postures. Fourty-two of these studies examined the impact of various postures on equine welfare, for example, by assessing behavior, physiological stress parameters, health or rider-horse interaction. Thirty-five studies examined the impact of various postures on gymnastics (e.g. kinematics, shifts in weight distribution, muscle activity, airway functioning or overall workload). For the meta-analysis a dataset containing information from each of the individual studies was created. Data included information such as type, degree, duration and circumstances of hyperflexion applied in that particular study as well as information on the horses (e.g., sport discipline, level of training, breed) and on the study design (e.g., size of study and experimental or epidemiological research design). The results of the study regarding the impact of hyperflexion on a) welfare and b) gymnastics were coded as positive (1), insignificant or contradictory (0) or negative (-1). The significant majority of studies (88%) concluded that a hyperflexed head and neck posture negatively impacts welfare. Just one study suggested welfare advantages of training in a hyperflexed head and neck posture. An analysis using a generalized linear mixed model to assess the influence of the above factors collated in the dataset revealed that none of these factors significantly influenced the probability of a study to detect negative welfare implications. Thus hyperflexing the neck appears to impair horses’ welfare regardless of, for example, the duration or the way of achieving hyperflexion. A concurrent assessment of the evidence for gymnastic benefits showed that approximately one quarter of studies conclude that there may be benefits, while another quarter of the studies conclude that hyperflexion has detrimental effects on gymnastics. Thus, on the costs-side there is a clear reduction in equine welfare and some undesirable gymnastic effects, as well as likely a compromised profile of the equestrian sports in public. Benefits, on the other hand, include some desirable gymnastic effects, and potentially increased control of the horse for the rider. On balance, it appears that the costs associated with hyperflexion exceed the potential benefits of the activity to both humans and horses.
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Baragli, P., Demuru, E., & Palagi, E. (2015). Mirror on the wall, who is the horsest of our all? Self-recognition in Equus caballus. In Proceedings of the 3. International Equine Science Meeting.
Abstract: Mirror Self-Recognition (MSR) is an extremely rare capacity in the animal kingdom that reveals the emergence of complex cognitive capacities (de Waal 2008). So far, MSR has been reported only in humans, chimpanzees (Gallup, 1970), bottlenose dolphins (Reiss and Marino, 2001) and Asian elephants (Plotnik et al, 2006), all species characterized by a highly developed cognition. There is growing evidence that domestic horses posses high cognitive abilities, such as cross-modal individual recognition (Proops et al, 2009), triadic post-conflict reunion to maintain social homeostasis (Cozzi et al, 2010), complex communicative systems (Whatan and McComb, 2014), flexibility in problem-solving (Lovrovich et al, 2015), and long-term memory (Hanggi and Ingersoll, 2009). All these capacities make horses a good candidate to test the ability of MSR in a domestic species. Through a classical MSR experimental paradigm (de Waal 2008) we tested eight horses living in social groups under semi-natural conditions (from the Italian Horse Protection rescue centre). Animals showing MSR typically go through four stages (Plotnik et al, 2006): (i) social response, (ii) physical mirror inspection (e.g., looking behind the mirror), (iii) repetitive mirror-testing behaviour (i.e., the beginning of mirror understanding), and (iv) self-directed behaviour (i.e., recognition of the mirror image as self). The final stage, known as the “mark-test”, is verified when a subject spontaneously uses the mirror to check for a coloured artificial mark on its own body which it cannot perceive otherwise. The horses underwent a three-phase “mark-test”: 1) with sham mark (transparent ultrasound water gel) positioned on both side at jaw level, 2) mark (yellow eye shadow mixed with ultrasound water gel) positioned on left side of jaw (with sham mark on the right), 3) mark (yellow eye shadow mixed with ultrasound water gel) positioned on right side of jaw (with sham mark on the left)
The mirror was one 0.5-cm-thick piece of 140x220-cm plexiglass glue on wood. Each test lasted one hour, horses were tested once a day, in consecutive days and at the same time. Our preliminary result on 1 horse shows some changes in self-directed behaviours which can be attributed to presence of the coloured mark. Firstly, the presence of the coloured mark significantly increased the frequency of scratching on both sides of the muzzle (p < 0.0001). The most intriguing result (p < 0.0001) comes from the comparison of the scratching rates directed towards the coloured mark side (N = 41) and the sham mark side (N = 23). Under the control condition (i.e. sham mark on both sides) no statistical difference was found for the scratching rates directed to the muzzle sides (dx N = 8; sx N = 5). Although further analyses are needed to confirm these preliminary results, our finding opens new scenarios about the evolution of Mirror Self-Recognition. The capacity of horses to recognize themselves in a mirror may be the outcome of an evolutionary convergence process driven by the cognitive pressures imposed by a complex social system and maintained despite thousands years of domestication.
Keywords:
Domestic horse · Mark test · Socio-cognitive skills · Self-awareness
References
De Waal FBM (2008) The thief in the mirror. PloS Biol 6(8):e201
Gallup GG Jr (1970) Chimpanzees: Self-recognition. Science 167: 86-87.
Reiss D, Marino L (2001). Mirror self-recognition in the bottlenose dolphin: A case of cognitive convergence. Proc Natl Acad Sci U S A 98:5937-5942.
Plotnik J, de Waal FBM, Reiss D (2006) Self-recognition in an Asian elephant. Proc Natl Acad Sci U S A 103: 17053-17057.
Proops L, McComb K, Reby D. (2009) Cross-modal individual recognition in domestic horses (Equus caballus). Proc Nat Acad Sci USA;106:947-951.
Cozzi A, Sighieri C, Gazzano A, Nicol CJ, Baragli P. Post-conflict friendly reunion in a permanent group of horses (Equus caballus). Behav Process 2010;85:185-190.
Wathan J, McComb K. The eyes and ears are visual indicators of attention in domestic horses. Curr Biol 2014;24(15): R677-R679.
Lovrovich P, Sighieri C, Baragli P (2015) Following human-given cues or not? Horses (Equus caballus) get smarter and change strategy in a delayed three choice task. Appl Anim Behav Sci, in press.
Hanggi EB, Ingersoll JF. (2009) Long-term memory for categories and concepts in horses (Equus caballus). Anim Cogn; 12:451-462.
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Hintze, S., Smith, S., Patt, A., Bachmann, I., & Würbel, H. (2015). What eye wrinkles in horses tell us about their emotional state. In Proceedings of the 3. International Equine Science Meeting.
Abstract: Wrinkles above the eye ball are common in domestic horses but may differ in number and shape both between and within individuals. They are caused by contraction of the inner eye brow raiser, and some people working with horses call them “worry wrinkles”, considering them to reflect emotional states. However, as yet no study has formally investigated the relationship between eye wrinkles and emotional state in horses.
The aim of the present study was to induce states of different emotional valence and to assess whether positive emotional states would reduce the expression of eye wrinkles while negative emotional states would increase it. Sixteen horses were confronted in a balanced order with two presumably positively and two negatively valenced situations each. Positive situations included anticipation of a food reward (FA) and petting (P), negative situations included food competition (FC) and waving a plastic bag (PB). Each situation lasted for 60s (TRT) and was preceded by a 60s control phase (CON). Throughout CON and TRT pictures of the eyes were taken, and for each horse four pictures per situation (FA, P, FC, PB) and phase (CON and TRT) were randomly selected (n = 512) and scored in random order and blind to treatment for six outcome variables: overall impression (qualitative), number, angle and markedness of eye wrinkles, presence of eye white, and shape of eye lid.
Data were analysed separately for the right and left eye using linear mixed effects models (angle, number), generalised linear mixed models (eye white, markedness), and ordered logistic regression (qualitative, shape of eye lid), with “situation” (FA, P, FC, PB), “phase” (CON, TRT) and their two-way interaction as fixed effects.
Expression of eye wrinkles did not vary consistently across “situation” and “phase”. Independent of phase, eye white appeared less frequently during P than during FA (z=-3.15, p=0.009), FC (z=-2.94, p=0.02), and PB (z=4.17, p<0.001) in the left eye and during PB (z=4.10, p 0.001) in the right eye. Similarly, wrinkles were less marked during P compared to the other situations in the left eye (FA: z=3.15, p=0.009; FC: z=-2.94, p=0.017; PB: z=4.17, p<0.001) and compared to PB in the right eye (z=4.10, p=0.001), while no differences between situations occurred in number of wrinkles, overall impression and shape of eye lid for both eyes. Consistent with our hypothesis, P induced relaxation of the underlying muscle in the right eye resulting in a wider angle compared to its control phase (interaction situation*phase: F3,10=3.71, p=0.055; post-hoc comparison: z=-3.57, p=0.009), while FC induced muscle contraction, resulting in a sharper angle in the left eye (interaction situation*phase: F3,11=6.57, p=0.011; z=3.73, p=0.005).
We conclude that emotional valence may affect characteristics of eye winkle expression in horses which might therefore be a promising indicator of horses’ emotional states, but further research is needed to validate the relevant outcome variables.
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