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Stomp, M.; d'Ingeo, S.; Henry, S.; Cousillas, H.; Hausberger, M. |
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Title |
Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model |
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Journal Article |
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2021 |
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Applied Animal Behaviour Science |
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236 |
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105271 |
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Laterality; Electroencephalography; Theta wave; Welfare; Horses |
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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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0168-1591 |
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Equine Behaviour @ team @ |
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6629 |
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Rørvang, M.V.; Nielsen, B.L.; McLean, A.N. |
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Title |
Sensory Abilities of Horses and Their Importance for Equitation Science |
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Journal Article |
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2020 |
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Frontiers in Veterinary Science |
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7 |
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633 |
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Vision, hearing, olfaction, taste, and touch comprise the sensory modalities of most vertebrates. With these senses, the animal receives information about its environment. How this information is organized, interpreted, and experienced is known as perception. The study of the sensory abilities of animals and their implications for behavior is central not only to ethology but also to animal welfare. Sensory ability, perception, and behavior are closely linked. Horses and humans share the five most common sensory modalities, however, their ranges and capacities differ, so that horses are unlikely to perceive their surroundings in a similar manner to humans. Understanding equine perceptual abilities and their differences is important when horses and human interact, as these abilities are pivotal for the response of the horse to any changes in its surroundings. This review aims to provide an overview of the current knowledge on the sensory abilities of horses. The information is discussed within an evolutionary context and also includes a practical perspective, outlining potential ways to mitigate risks of injuries and enhance positive horse-human interactions. The equine sensory apparatus includes panoramic visual capacities with acuities similar to those of red-green color-blind humans as well as aural abilities that, in some respects exceed human hearing and a highly developed sense of smell, all of which influence how horses react in various situations. Equine sensitivity to touch has been studied surprisingly sparingly despite tactile stimulation being the major interface of horse training. We discuss the potential use of sensory enrichment/positive sensory stimulation to improve the welfare of horses in various situations e.g. using odors, touch or sound to enrich the environment or to appease horses. In addition, equine perception is affected by factors such as breed, individuality, age, and in some cases even color, emphasizing that different horses may need different types of management. Understanding the sensory abilities of horses is central to the emerging discipline of equitation science, which comprises the gamut of horse-human interactions. Therefore, sensory abilities continue to warrant scientific focus, with more research to enable us to understand different horses and their various needs. |
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2297-1769 |
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Equine Behaviour @ team @ |
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6634 |
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Ahrendt, L.P.; Labouriau, R.; Malmkvist, J.; Nicol, C.J.; Christensen, J.W. |
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Development of a standard test to assess negative reinforcement learning in horses |
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2015 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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169 |
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38-42 |
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Algometry; Horse behaviour; Learning performance; Operant conditioning; Pressure-release; Horse training |
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Most horses are trained by negative reinforcement. Currently, however, no standardised test for evaluating horses' negative reinforcement learning ability is available. The aim of this study was to develop an objective test to investigate negative reinforcement learning in horses. Twenty-four Icelandic horses (3 years old) were included in this study. The horses were tested in a pressure-release task on three separate days with 10, 7 and 5 trials on each side, respectively. Each trial consisted of pressure being applied on the hindquarter with an algometer. The force of the pressure was increased until the horse moved laterally away from the point of pressure. There was a significant decrease in required force over trials on the first test day (P<0.001), but not the second and third day. The intercepts on days 2 and 3 differed significantly from day 1 (P<0.001), but not each other. Significantly stronger force was required on the right side compared to the left (P<0.001), but there was no difference between first and second side tested (P=0.56). Individual performance was evaluated by median-force and the change in force over trials on the first test day. These two measures may explain different characteristics of negative reinforcement learning. In conclusion, this study presents a novel, standardised test for evaluating negative reinforcement learning ability in horses. |
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0168-1591 |
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Equine Behaviour @ team @ |
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6650 |
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Mejdell, C.M.; Buvik, T.; Jørgensen, G.H.M.; Bøe, K.E. |
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Horses can learn to use symbols to communicate their preferences |
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2016 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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184 |
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66-73 |
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Operant conditioning; Blanket; Rug; Thermoregulation; Cognition; Clicker training |
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This paper describes a method in which horses learn to communicate by touching different neutral visual symbols, in order to tell the handler whether they want to have a blanket on or not. Horses were trained for 10-15min per day, following a training program comprising ten steps in a strategic order. Reward based operant conditioning was used to teach horses to approach and touch a board, and to understand the meaning of three different symbols. Heat and cold challenges were performed to help learning and to check level of understanding. At certain stages, a learning criterion of correct responses for 8-14 successive trials had to be achieved before proceeding. After introducing the free choice situation, on average at training day 11, the horse could choose between a “no change” symbol and the symbol for either “blanket on” or “blanket off” depending on whether the horse already wore a blanket or not. A cut off point for performance or non-performance was set to day 14, and 23/23 horses successfully learned the task within this limit. Horses of warm-blood type needed fewer training days to reach criterion than cold-bloods (P<0.05). Horses were then tested under differing weather conditions. Results show that choices made, i.e. the symbol touched, was not random but dependent on weather. Horses chose to stay without a blanket in nice weather, and they chose to have a blanket on when the weather was wet, windy and cold (χ2=36.67, P<0.005). This indicates that horses both had an understanding of the consequence of their choice on own thermal comfort, and that they successfully had learned to communicate their preference by using the symbols. The method represents a novel tool for studying preferences in horses. |
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0168-1591 |
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Equine Behaviour @ team @ |
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6651 |
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Suagee-Bedore, J.K.; Linden, D.R.; Bennett-Wimbush, K. |
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Effect of Pen Size on Stress Responses of Stall-Housed Horses Receiving One Hour of Daily Turnout |
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Journal Article |
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2021 |
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Journal of Equine Veterinary Science |
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J. Equine Vet. Sci. |
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98 |
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103366 |
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Agonistic behaviors; Cortisol; Group turnout; Paddock sizes |
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Group turnout provides important socializing opportunities for horses, particularly those that are primarily stalled. A high percentage of equine injuries occur during group turnout, which could partly be due to the physical constraints of fencing. To investigate appropriate paddock sizes for group turnouts, horses (n = 12) from a single herd were divided into groups of 4, stalled for 24 hours, and then turned out for 1 hour into one of three differently sized pens: 342, 263, and 184 m2 per horse. Groups rotated through pens across 3 days, receiving one treatment per day. Blood was sampled for cortisol concentrations at 08:00 hours each morning, and then at 15 and 60 minutes into the turn out sessions, and 60 minutes after return to individual stalls. Groups rotated through three turnout times: 09:00, 12:00, 14:00 hours. Counts of agonistic behaviors (chasing, contact biting, and kicking) and low-level threats (pinned ears, tail swishing, bite and kick threats) were recorded. When turned out in pens that provided 342 m2 per horse, horses exhibited reduced plasma cortisol concentrations by 15 minutes after turnout and at 1 hour after return to their stalls (P < .05). Horses in pens providing 184 m2 per horse exhibited greater agonistic (P < .001) and low-level threat (P < .01) behaviors than horses in larger pens. These data provide insight into appropriate pen sizes for horses from established herds. Providing at least 342 m2 per horse may reduce the chance of injury in horses accustomed to group turnout. |
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0737-0806 |
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Equine Behaviour @ team @ |
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6694 |
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Christensen, J.W.; Munk, R.; Hawson, L.; Palme, R.; Larsen, T.; Egenvall, A.; König von Borstel, U.U.; Rørvang, M.V. |
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Rider effects on horses' conflict behaviour, rein tension, physiological measures and rideability scores |
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Journal Article |
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2021 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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234 |
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105184 |
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Equitation science; Heart rate; Horse riding; Sport horse; Temperament; Training cues |
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Many breeding organisations include a subjective scoring of rideability by a professional rider into their evaluation of sports horses, but the consistency and reliability of the scoring system is debateable. The aim of this study was to investigate (i) whether professional riders agree in their scoring of rideability, and (ii) whether rideability scores are affected by rein tension, horse conflict behaviour, heart rate, and salivary cortisol, and (iii) whether riders induce different levels of conflict behaviour and physiological responses in the horses. Ten professional, female riders each rode 10 dressage horses (level M German scale; n = 100 combinations) through a standardised dressage test (10 min warm-up followed by a 4-min test) and subsequently scored the horses for rideability on the official 1-10 scale (1 = poor to 10 = excellent) from the Danish Riding Federation. Rein tension, horse heart rate, saliva cortisol and conflict behaviour were measured for each rider-horse pair. The riders were inconsistent in their scoring of rideability to the individual horses, e.g. scores for one of the horses ranged from 1 to 8. There was a significant effect of rider (P = 0.003) and the frequency of conflict behaviour (undesired head movements: P < 0.001, breaking the gait: P = 0.013, and other evasive behaviour: P = 0.032) on rideability scores, i.e. the more conflict behaviour the lower the score. There was no significant effect of rein tension and the physiological measures on rideability scores. However, there was a significant effect of rider on rein tension, horses' heart rate and increases in saliva cortisol concentrations and a tendency for some types of conflict behaviour, suggesting that some riders induced more discomfort in the horses. Future studies could help shed light on which elements of riding style are particularly important for sports horse welfare. In conclusion, this study found a large variation in rideability scores assigned to ten sports horses by ten professional riders. Rideability scores were dependent on the level of horse conflict behaviour, but not rein tension and physiological measures. Further studies are needed to improve the objectivity, consistency and reliability of rideability assessment of sports horses. |
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0168-1591 |
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Equine Behaviour @ team @ |
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6696 |
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Goursot, C.; Düpjan, S.; Puppe, B.; Leliveld, L.M.C. |
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Affective styles and emotional lateralization: A promising framework for animal welfare research |
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2021 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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237 |
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105279 |
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Individuality; Motor lateralization; Hemispheric dominance; Motivational tendencies; Emotional reactivity; Emotional regulation |
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The growing recognition of animals as individuals has broader implications for farm animal welfare research. Even under highly standardized on-farm conditions, farm animals show heterogeneous but individually consistent behavioural patterns towards various stimuli, based on how they appraise these stimuli. As a result, animal welfare is likely to be highly individual as well, and studying the proximate mechanisms underlying distinct individual behaviour patterns and appraisal will improve animal welfare research. We propose to extend the framework of affective styles to bridge the gap between existing research fields on animal personality and affective states. Affective styles refer to consistent individual differences in emotional reactivity and regulation and can be predicted by baseline cerebral lateralization. Likewise, animals with consistent left or right motor biases--a proxy measure of individual patterns in cerebral lateralization--have been shown to differ in their personality, emotional reactivity, motivational tendencies or coping styles. In this paper, we present the current knowledge of the links between laterality and stable individual traits in behaviour and affect in light of hypotheses on emotional lateralization. Within our suggested framework, we make recommendations on how to investigate affective styles in non-human animals and give practical examples. This approach has the potential to promote a science of affective styles in nonhuman animals and significantly advance research on animal welfare. |
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Equine Behaviour @ team @ |
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6698 |
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Leliveld, L.M.C.; Düpjan, S.; Tuchscherer, A.; Puppe, B. |
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Hemispheric Specialization for Processing the Communicative and Emotional Content of Vocal Communication in a Social Mammal, the Domestic Pig |
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2020 |
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Frontiers in Behavioral Neuroscience |
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Front. Behav. Neurosci. |
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14 |
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596758 |
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In humans, speech perception is lateralized, with the left hemisphere of the brain dominant in processing the communicative content and the right hemisphere dominant in processing the emotional content. However, still little is known about such a division of tasks in other species. We therefore investigated lateralized processing of communicative and emotionally relevant calls in a social mammal, the pig (Sus scrofa). Based on the contralateral connection between ears and hemispheres, we compared the behavioural and cardiac responses of 36 young male pigs during binaural and monaural (left or right) playback to the same sounds. The playback stimuli were calls of social isolation and physical restraint, whose communicative and emotional relevance, respectively, were validated prior to the test by acoustic analyses and during binaural playbacks. There were indications of lateralized processing mainly in the initial detection (left head-turn bias, indicating right hemispheric dominance) of the more emotionally relevant restraint calls. Conversely, there were indications of lateralized processing only in the appraisal (increased attention during playback to the right ear) of the more communicative relevant isolation calls. This implies differential involvement of the hemispheres in the auditory processing of vocalizations in pigs and thereby hints at similarities in the auditory processing of vocal communication in non-human animals and speech in humans. Therefore, these findings provide interesting new insight in the evolution of human language and auditory lateralization. |
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Equine Behaviour @ team @ |
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6699 |
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Reeve, H. Kern |
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Evolutionarily stable communication between kin: a general model |
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1997 |
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Proceedings of the Royal Society B: Biological Sciences |
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Proc. Roy. Soc. Lond. B Biol. Sci. |
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1037-1040. |
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Signalling Systems |
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At present, the most general evolutionary theory of honest communication is Grafen's model of Zahavi's 'handicap' signalling system, in which honesty of signals about the signaller's quality (e.g. mate suitability or fighting ability) is maintained by the differentially high cost of signals to signallers having lower quality. The latter model is here further generalized to include any communication between signallers and receivers that are genetically related (e.g. parents and begging offspring, cooperative or competing siblings). Signalling systems involving relatives are shown to be evolutionarily stable, despite a potential pay-off for false signalling, if the Zahavian assumption of differential signal costs holds and there are diminishing reproductive returns to the signaller as the receiver's assessed value of its attribute increases, or if, regardless of whether the Zahavian assumption holds, signallers with high values of the attribute benefit more from a given receiver assessment than signallers with low values (e.g. begging chicks that are hungrier benefit more from being fed). In stable systems of signalling among kin, it is also shown to be generally true that (i) levels of signalling and thus observed signal costs will decline as relatedness increases or as the receiver's reproductive penalty for erroneous assessment increases, and (ii) receivers will consistently, altruistically overestimate the true value of the signalled attribute. |
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refbase @ user @ |
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Hartmann, E.; Søndergaard, E.; Keeling, L.J. |
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Keeping horses in groups: A review |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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Equine; Behaviour; Welfare; Housing; Social |
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Although husbandry conditions for horses have improved over the last decades, many horses are still kept singly with limited or no physical contact to other horses. This is surprising, given the fact that keeping horses in groups is recognised best to fulfil their physical and behavioural needs, especially their need for social contact with conspecifics, as well as to have a beneficial effect on horse–human interactions during training. Group housing of farm animals is widely applied in practice. As a consequence, scientists have investigated numerous aspects of group housing to help further improve animal welfare and human–animal interactions under these conditions. However, compared to this literature available in farm animals, and the plentiful studies conducted of feral horse populations, there is much less done when it comes to the management of horses kept in groups in the domestic environment. In particular, limited scientific information is available into the effect of group size and group composition on behaviour and methods of introducing new horses into established groups, even though problems related to social integration are repeatedly taken as arguments against keeping horses in groups. This review, therefore, aims to provide an overview of the current scientific knowledge regarding keeping horses in groups. Furthermore, it aims to give insight into whether or not some of the concerns related to keeping horses in groups are justified and to review scientifically based solutions that could be useful in practice to improve horse welfare and human safety. |
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0168-1591 |
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Call Number |
Equine Behaviour @ team @ |
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5490 |
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