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Outram, A. K., Stear, N. A., Bendrey, R., Olsen, S., Kasparov, A., Zaibert, V., et al. (2009). The Earliest Horse Harnessing and Milking. Science, 323(5919), 1332–1335.
Abstract: Horse domestication revolutionized transport, communications, and warfare in prehistory, yet the identification of early domestication processes has been problematic. Here, we present three independent lines of evidence demonstrating domestication in the Eneolithic Botai Culture of Kazakhstan, dating to about 3500 B.C.E. Metrical analysis of horse metacarpals shows that Botai horses resemble Bronze Age domestic horses rather than Paleolithic wild horses from the same region. Pathological characteristics indicate that some Botai horses were bridled, perhaps ridden. Organic residue analysis, using δ13C and δD values of fatty acids, reveals processing of mare's milk and carcass products in ceramics, indicating a developed domestic economy encompassing secondary products.
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Imbert, C., Caniglia, R., Fabbri, E., Milanesi, P., Randi, E., Serafini, M., et al. (2016). Why do wolves eat livestock?: Factors influencing wolf diet in northern Italy. Biological Conservation, 195, 156–168.
Abstract: Thanks to protection by law and increasing habitat restoration, wolves (Canis lupus) are currently re-colonizing Europe from the surviving populations of Russia, the Balkan countries, Spain and Italy, raising the need to update conservation strategies. A major conservation issue is to restore connections and gene flow among fragmented populations, thus contrasting the deleterious consequences of isolation. Wolves in Italy are expanding from the Apennines towards the Alps, crossing the Ligurian Mountains (northern Italy) and establishing connections with the Dinaric populations. Wolf expansion is threatened by poaching and incidental killings, mainly due to livestock depredations and conflicts with shepherds, which could limit the establishment of stable populations. Aiming to find out the factors affecting the use of livestock by wolves, in this study we determined the composition of wolf diet in Liguria. We examined 1457 scats collected from 2008 to 2013. Individual scats were genotyped using a non-invasive genetic procedure, and their content was determined using microscopical analyses. Wolves in Liguria consumed mainly wild ungulates (64.4%; in particular wild boar Sus scrofa and roe deer Capreolus capreolus) and, to a lesser extent, livestock (26.3%; in particular goats Capra hircus). We modeled the consumption of livestock using environmental features, wild ungulate community diversity, husbandry characteristics and wolf social organization (stable packs or dispersing individuals). Wolf diet varied according to years and seasons with an overall decrease of livestock and an increase of wild ungulate consumption, but also between packs and dispersing individuals with greater livestock consumption for the latter. The presence of stable packs, instead of dispersing wolves, the adoption of prevention measures on pastures, roe deer abundance, and the percentage of deciduous woods, reduced predation on livestock. Thus, we suggest promoting wild ungulate expansion, the use of prevention tools in pastures, and supporting wolf pack establishment, avoiding lethal control and poaching, to mitigate conflicts between wolf conservation and husbandry.
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López-Bao, J. V., Sazatornil, V., Llaneza, L., & Rodríguez, A. (2013). Indirect Effects on Heathland Conservation and Wolf Persistence of Contradictory Policies that Threaten Traditional Free-Ranging Horse Husbandry. Conservation Letters, 6(6), 448–455.
Abstract: Abstract Conservation agencies within the European Union promote the restoration of traditional land uses as a cost-effective way to preserve biodiversity outside reserves. Although the European Union pursues the integration of the environment into strategic decision-making, it also dictates sectoral policies that may damage farmland biodiversity. We illustrate this point by outlining the socioeconomic factors that allow the persistence of traditional free-ranging horse husbandry in Galicia, northwestern Spain. Free-ranging Galician mountain ponies provide ecological and socioeconomic services including the prevention of forest fires, the maintenance of heathlands and wolves, and the attenuation of wolf-human conflicts. This traditional livestock system may have persisted because it entails negligible costs for farmers. Wolf predation upon Galician mountain ponies does not threaten farmer's economies and seems to be tolerated better than attacks to more valuable stock. Recently, European Union's regulations on animal welfare, carcass management, or meat production put new economic and administrative burdens on farmers, make free-ranging horse rearing economically unsustainable, and incentivize its abandonment. The aim of the European Union to integrate environmental policies may be successful to preserve farmland biodiversity only through careful anticipation of the side effects of apparently unrelated regulations on the fragile equilibrium that sustain traditional land uses.
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Lansade, L., Colson, V., Parias, C., Trösch, M., Reigner, F., & Calandreau, L. (2020). Female horses spontaneously identify a photograph of their keeper, last seen six months previously. Scientific Reports, 10(1), 6302.
Abstract: Horses are capable of identifying individual conspecifics based on olfactory, auditory or visual cues. However, this raises the questions of their ability to recognize human beings and on the basis of what cues. This study investigated whether horses could differentiate between a familiar and unfamiliar human from photographs of faces. Eleven horses were trained on a discrimination task using a computer-controlled screen, on which two photographs were presented simultaneously (32 trials/session): touching one was rewarded (S+) and the other not (S-). In the training phase, the S+ faces were of four unfamiliar people which gradually became familiar over the trials. The S- faces were novel for each trial. After the training phase, the faces of the horses' keepers were presented opposite novel faces to test whether the horses could identify the former spontaneously. A reward was given whichever face was touched to avoid any possible learning effect. Horses touched the faces of keepers significantly more than chance, whether it was their current keeper or one they had not seen for six months (t = 3.65; p < 0.004 and t = 6.24; p < 0.0001). Overall, these results show that horses have advanced human face-recognition abilities and a long-term memory of those human faces.
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Janczarek, I., Stachurska, A., Kedzierski, W., Wisniewska, A., Ryzak, M., & Koziol, A. (2020). The intensity of physiological and behavioral responses of horses to predator vocalizations. BMC Veterinary Research, 16(1), 431.
Abstract: Predatory attacks on horses can become a problem in some parts of the world, particularly when considering the recovering gray wolf populations. The issue studied was whether horses transformed by humans and placed in stable-pasture environments had retained their natural abilities to respond to predation risk. The objective of the study was to determine the changes in cardiac activity, cortisol concentrations, and behavior of horses in response to the vocalizations of two predators: the gray wolf (Canis lupus), which the horses of the breed studied had coevolved with but not been exposed to recently, and Arabian leopard (Panthera pardus nimr), from which the horses had been mostly isolated. In addition, we hypothesized that a higher proportion of Thoroughbred (TB) horse ancestry in the pedigree would result in higher emotional excitability in response to predator vocalizations. Nineteen horses were divided into groups of 75%, 50% and 25% TB ancestry. The auditory test conducted in a paddock comprised a 10-min prestimulus period, a 5-min stimulus period when one of the predators was heard, and a 10-min poststimulus period without any experimental stimuli.
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Scopa, C., Greco, A., Contalbrigo, L., Fratini, E., Lanatà, A., Scilingo, E. P., et al. (2020). Inside the Interaction: Contact With Familiar Humans Modulates Heart Rate Variability in Horses. Frontiers in Veterinary Science, 7, 998.
Abstract: A human-animal relationship can be developed through subsequent interactions, affected by the positive or negative emotional valence of the proceeding one. Horses implement a process of categorization to classify humans with whom they interact as positive, negative, or neutral stimuli by evaluating the kind of approach and the nature of the contact. In these terms, human-animal interactions are emotionally charged events, eliciting specific emotional states in both subjects involved. Although the human-horse relationship has been mainly investigated through behavioral analysis, physiological indicators are needed for a more objective assessment of the emotional responses. Heart rate variability (HRV) is a commonly used autonomic nervous system (ANS) correlate estimating the sympathovagal balance as a psychophysiological marker of emotion regulation in horses. We have assumed that long-term positive relationships with humans may have a positive and immediate impact on the emotional arousal of the horse, detectable, via ANS activity, during the interaction. We analyzed horses' heartbeat dynamics during their interaction with either familiar or unfamiliar handlers, applying a standardized experimental protocol consisting of three different conditions shifting from the absence of interaction to physical contact. The ANS signals were monitored through an innovative non-invasive wearable system, not interfering with the unconscious emotional response of the animal. We demonstrated that horses appeared to feel more relaxed while physically interacting (e.g., grooming on the right side) with some familiar handlers compared to the same task performed by someone unfamiliar. The shift of the sympathovagal balance toward a vagal predominance suggests that the horses experienced a decrease in stress response as a function not only of the handler's familiarity but also of the type of interaction they are experiencing. These results constitute the objective evidence of horses' capacity to individually recognize a familiar person, adding the crucial role of familiarity with the handler as a paramount component of human-animal interaction. Our rigorous methodological approach may provide a significant contribution to various fields such as animal welfare while further investigating the emotional side of the human-animal relationships.
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Werhahn, H., Hessel, E. F., & Van den Weghe, H. F. A. (2012). Competition Horses Housed in Single Stalls (II): Effects of Free Exercise on the Behavior in the Stable, the Behavior during Training, and the Degree of Stress. Journal of Equine Veterinary Science, 32(1), 22–31.
Abstract: Although housing horses in single stalls limits their natural behavior to a great extent, this housing system is widespread in Germany, especially for competition horses. To improve the welfare of this system, free exercise on pastures or paddocks is deemed suitable, but it is also feared because of injuries and decreased willingness or motivation to perform. In the present study, three treatments were investigated with regard to their effect on the behavior of six competition horses in the stable, behavior during training, and on their degree of stress: daily training without free exercise (no turnout [NT]), solitary turnout for 2 hours after training, and 2-hour turnout in groups of two after training (group turnout). The horses' behavior in the stable was continuously analyzed through video recordings (2 pm to 6 am) on 3 days at the end of each treatment. The degree of stress was evaluated daily by heart rate variability at rest. The behavior during training was evaluated by a questionnaire answered by the riders, and the distance covered during training was measured by global positioning system. When NT was allowed, the horses showed less lying in the stable compared with the treatments with turnout. Heart rate variability measurements resulted in great individual differences, but generally, there was a higher degree of stress shown with the treatment NT according to the following parameters: standard deviation of inter-beat-intervals (SDNN), square root of the mean of the sum of the squares of differences between successive inter-beat-intervals (RMSSD), and ratio between low frequency and high frequency (LF/HF). The willingness to perform was evaluated as being slightly better in the treatments with turnout than in the treatment without turnout.
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McBride, S. D., Parker, M. O., Roberts, K., & Hemmings, A. (2017). Applied neurophysiology of the horse; implications for training, husbandry and welfare. Appl. Anim. Behav. Sci., 190, 90–101.
Abstract: Understanding the neural circuits underlying equine behaviour has the potential to help optimise strategies of husbandry and training. This review discusses two areas of neurophysiological research in a range of species and relates this information to the horse. The first discussion focuses on mechanisms of learning and motivation and assesses how this information can be applied to improve the training of the horse. The second concerns the identification of the equine neurophysiological phenotype, through behavioural and genetic probes, as a way of improving strategies for optimal equine husbandry and training success. The review finishes by identifying directions for future research with an emphasis on how neurophysiological systems (and thus behaviour) can be modified through strategic husbandry. This review highlights how a neurophysioloigical understanding of horse behaviour can play an important role in attaining the primary objectives of equitation science as well as improving the welfare of the horse.
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Stomp, M., d'Ingeo, S., Henry, S., Cousillas, H., & Hausberger, M. (2021). Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model. Appl. Anim. Behav. Sci., 236, 105271.
Abstract: Assessing the animal welfare state is a challenge given the subjective individual cognitive and emotional processing involved. Electroencephalography (EEG) spectrum analysis has proved an ecologically valid recording situation to assess the link between brain processes and affective or cognitive states in humans: a higher slow wave/fast wave ratio has been associated with a positive internal state. In particular, a high production of theta power (3-8 Hz) has been related to positive emotions. On the other hand, it has been hypothesized that a left hemisphere (LH) dominance may be associated with a better welfare state. Here, we test the hypothesis that individual differences in the resting-state quantitative EEG power spectrum of adult horses (N = 18) and its lateralization pattern may reflect individual subjective perception of their conditions of life and welfare state. The results show clear individual differences in the proportions of the different waves and their inter-hemispheric distribution. Three different EEG power spectrum profiles were highlighted, from a bilateral predominance of theta waves in horses in a more positive welfare state to a bilateral predominance of beta waves in horses with clear expressions of compromised welfare. Interestingly, particular correlations were found between wave power activity and welfare parameters. We found a negative correlation between the number of stereotypic behaviours per hour and the median proportion of theta waves in the left hemisphere. and between the overall state (total chronic stress score) of welfare and gamma production in the right hemisphere (RH). These findings go along the hypothesis of a particular involvement of the left hemisphere for positive processing and of the right hemisphere for negative processing. However, the pattern of laterality did not appear as the most important feature here as both extreme clusters in terms of welfare showed bilateral predominance of one wave type. It is possible that hemispheric specialization makes more sense during acute emotion-inducing conditions rather than in this resting-state context (i.e. in absence of any high emotion-inducing stimulation), although the opposition gamma versus theta waves between both hemispheres in the horses with an intermediate welfare state is noticeable and intriguing. It seems that bilateral but also LH theta activity is a promising neurophysiological marker of good welfare in horses, while a bilateral or RH high production of gamma waves should alert about potential welfare alterations. Quantitative resting-state EEG power spectrum appears as a highly promising tool for exploring the brain processes involved in the subjective perception of chronic welfare, as a useful complementary tool for welfare assessment.
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Stomp, M., d'Ingeo, S., Henry, S., Cousillas, H., & Hausberger, M. (2021). Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model. Applied Animal Behaviour Science, 236, 105271.
Abstract: Assessing the animal welfare state is a challenge given the subjective individual cognitive and emotional processing involved. Electroencephalography (EEG) spectrum analysis has proved an ecologically valid recording situation to assess the link between brain processes and affective or cognitive states in humans: a higher slow wave/fast wave ratio has been associated with a positive internal state. In particular, a high production of theta power (3-8 Hz) has been related to positive emotions. On the other hand, it has been hypothesized that a left hemisphere (LH) dominance may be associated with a better welfare state. Here, we test the hypothesis that individual differences in the resting-state quantitative EEG power spectrum of adult horses (N = 18) and its lateralization pattern may reflect individual subjective perception of their conditions of life and welfare state. The results show clear individual differences in the proportions of the different waves and their inter-hemispheric distribution. Three different EEG power spectrum profiles were highlighted, from a bilateral predominance of theta waves in horses in a more positive welfare state to a bilateral predominance of beta waves in horses with clear expressions of compromised welfare. Interestingly, particular correlations were found between wave power activity and welfare parameters. We found a negative correlation between the number of stereotypic behaviours per hour and the median proportion of theta waves in the left hemisphere. and between the overall state (total chronic stress score) of welfare and gamma production in the right hemisphere (RH). These findings go along the hypothesis of a particular involvement of the left hemisphere for positive processing and of the right hemisphere for negative processing. However, the pattern of laterality did not appear as the most important feature here as both extreme clusters in terms of welfare showed bilateral predominance of one wave type. It is possible that hemispheric specialization makes more sense during acute emotion-inducing conditions rather than in this resting-state context (i.e. in absence of any high emotion-inducing stimulation), although the opposition gamma versus theta waves between both hemispheres in the horses with an intermediate welfare state is noticeable and intriguing. It seems that bilateral but also LH theta activity is a promising neurophysiological marker of good welfare in horses, while a bilateral or RH high production of gamma waves should alert about potential welfare alterations. Quantitative resting-state EEG power spectrum appears as a highly promising tool for exploring the brain processes involved in the subjective perception of chronic welfare, as a useful complementary tool for welfare assessment.
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