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Weissing, F. J. (2011). Animal behaviour: Born leaders. Nature, 474(7351), 288–289.
Abstract: Social animals face a dilemma. To reap the benefits of group living, they have to stay together. However, individuals differ in their preferences as to where to go and what to do next. If all individuals follow their own preferences, group coherence is undermined, resulting in an outcome that is unfavourable for everyone. Neglecting one's own preferences and following a leader is one way to resolve this coordination problem. But what attributes make an individual a 'leader'? A modelling study by Johnstone and Manica1 illuminates this question.
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Dyer, J. R. G., Johansson, A., Helbing, D., Couzin, I. D., & Krause, J. (2009). Leadership, consensus decision making and collective behaviour in humans. Phil. Trans. Biol. Sci., 364(1518), 781–789.
Abstract: This paper reviews the literature on leadership in vertebrate groups, including recent work on human groups, before presenting the results of three new experiments looking at leadership and decision making in small and large human groups. In experiment 1, we find that both group size and the presence of uninformed individuals can affect the speed with which small human groups (eight people) decide between two opposing directional preferences and the likelihood of the group splitting. In experiment 2, we show that the spatial positioning of informed individuals within small human groups (10 people) can affect the speed and accuracy of group motion. We find that having a mixture of leaders positioned in the centre and on the edge of a group increases the speed and accuracy with which the group reaches their target. In experiment 3, we use large human crowds (100 and 200 people) to demonstrate that the trends observed from earlier work using small human groups can be applied to larger crowds. We find that only a small minority of informed individuals is needed to guide a large uninformed group. These studies build upon important theoretical and empirical work on leadership and decision making in animal groups.
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Broad, K. D., Curley, J. P., & Keverne, E. B. (2006). Mother-infant bonding and the evolution of mammalian social relationships. Phil. Trans. Biol. Sci., 361(1476), 2199–2214.
Abstract: A wide variety of maternal, social and sexual bonding strategies have been described across mammalian species, including humans. Many of the neural and hormonal mechanisms that underpin the formation and maintenance of these bonds demonstrate a considerable degree of evolutionary conservation across a representative range of these species. However, there is also a considerable degree of diversity in both the way these mechanisms are activated and in the behavioural responses that result. In the majority of small-brained mammals (including rodents), the formation of a maternal or partner preference bond requires individual recognition by olfactory cues, activation of neural mechanisms concerned with social reward by these cues and gender-specific hormonal priming for behavioural output. With the evolutionary increase of neocortex seen in monkeys and apes, there has been a corresponding increase in the complexity of social relationships and bonding strategies together with a significant redundancy in hormonal priming for motivated behaviour. Olfactory recognition and olfactory inputs to areas of the brain concerned with social reward are downregulated and recognition is based on integration of multimodal sensory cues requiring an expanded neocortex, particularly the association cortex. This emancipation from olfactory and hormonal determinants of bonding has been succeeded by the increased importance of social learning that is necessitated by living in a complex social world and, especially in humans, a world that is dominated by cultural inheritance. © 2006 The Royal Society.
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Garamszegi, L. Z., Møller, A. P., & Erritzøe, J. (2002). Coevolving avian eye size and brain size in relation to prey capture and nocturnality. Proc Roy Soc Lond B Biol Sci, 269(1494), 961–967.
Abstract: Behavioural adaptation to ecological conditions can lead to brain size evolution. Structures involved in behavioural visual information processing are expected to coevolve with enlargement of the brain. Because birds are mainly vision–oriented animals, we tested the predictions that adaptation to different foraging constraints can result in eye size evolution, and that species with large eyes have evolved large brains to cope with the increased amount of visual input. Using a comparative approach, we investigated the relationship between eye size and brain size, and the effect of prey capture technique and nocturnality on these traits. After controlling for allometric effects, there was a significant, positive correlation between relative brain size and relative eye size. Variation in relative eye and brain size were significantly and positively related to prey capture technique and nocturnality when a potentially confounding variable, aquatic feeding, was controlled statistically in multiple regression of independent linear contrasts. Applying a less robust, brunching approach, these patterns also emerged, with the exception that relative brain size did not vary with prey capture technique. Our findings suggest that relative eye size and brain size have coevolved in birds in response to nocturnal activity and, at least partly, to capture of mobile prey.
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Lesimple, C., Reverchon-Billot, L., Galloux, P., Stomp, M., Boichot, L., Coste, C., et al. (2020). Free movement: A key for welfare improvement in sport horses? Appl. Anim. Behav. Sci., 225, 104972.
Abstract: Horses, and in particular sport horses, remain housed predominantly in single stalls. One of the main reported reasons is the fear that they will become agitated and injure themselves and thereby impair their performance if released in paddocks. The hour spent daily at work is also assumed to be sufficient to satisfy the horses' needs for locomotion. Growing scientific evidence shows that single stall housing has negative consequences on horses' welfare and that time for free movement is necessary. Our aim was to assess the feasibility of allowing sport horses used to staying permanently in their stall (except for 1 h riding/day) daily free time in a paddock and to evaluate its potential impact on their welfare using two behavioural reliable indicators (stereotypic behaviours and ear position), as well as selected blood parameters (blood cell count, oxytocin and serotonin concentrations). Two experiments were conducted on the same site. The first experiment evaluated sport horses' habituation to the novel situation of daily sessions in a paddock, and recorded welfare indicators in their stall before and during the experiment, comparing horses that were taken outdoors (experimental) and those that stayed in their stall (control). The second experiment evaluated the impact of this daily free time in a paddock on horses' welfare and its durability, focusing on positive indicators. Behavioural observations in paddocks showed that even horses that had never experienced free movement outside their stall habituated rapidly to this situation. The presence of hay in the paddock, may have speeded up habituation. Their restricted living conditions were associated with abnormalities in blood cell count that were not overcome during the time of daily paddock sessions but behavioural indicators showed that their welfare improved. In the second study, the experimental horses' welfare improved during the paddock release period, in particular their stereotypic behaviours decreased and oxytocin levels increased. No effects on serotonin concentrations could be evidenced. These effects were directly associated with being in paddock, as the indicators returned rapidly to their previous levels indicating compromised welfare when the paddock release sessions stopped. In conclusion, it can be recommended to release sport horses for free movement in paddocks as welfare is improved and subjective assessment by caretakers indicated minimal risks.
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Siniscalchi, M., Sasso, R., Pepe, A. M., Vallortigara, G., & Quaranta, A. (2010). Dogs turn left to emotional stimuli. Behav. Brain. Res., 208(2), 516–521.
Abstract: During feeding behaviour, dogs were suddenly presented with 2D stimuli depicting the silhouette of a dog, a cat or a snake simultaneously into the left and right visual hemifields. A bias to turn the head towards the left rather than the right side was observed with the cat and snake stimulus but not with the dog stimulus. Latencies to react following stimulus presentation were lower for left than for right head turning, whereas times needed to resume feeding behaviour were higher after left rather than after right head turning. When stimuli were presented only to the left or right visual hemifields, dogs proved to be more responsive to left side presentation, irrespective of the type of stimulus. However, cat and snake stimuli produced shorter latencies to react and longer times to resume feeding following left rather than right monocular visual hemifield presentation. Results demonstrate striking lateralization in dogs, with the right side of the brain more responsive to threatening and alarming stimuli. Possible implications for animal welfare are discussed.
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Munsters, C. C. B. M., Visser, K. E. K., van den Broek, J., & Sloet van Oldruitenborgh-Oosterbaan, M. M. (2012). The influence of challenging objects and horse-rider matching on heart rate, heart rate variability and behavioural score in riding horses. The Veterinary Journal, 192(1), 75–80.
Abstract: A good horse-rider ‘match’ is important in the context of equine welfare. To quantify the influence of repetition and horse-rider matching on the stress of horses encountering challenging objects, 16 Warmblood horses were ridden in a test-setting on three occasions. On each occasion the horse was ridden by a different rider and was challenged by three objects (A–C). Heart rate (HR), heart rate variability (HRV) of horse and rider, and behaviour score (BS) of the horse were obtained for each object and as a total for each test. The horse-rider interaction was evaluated with each combination and assessed as ‘matching’ or ‘mismatching’, and the horses were categorised as ‘compliant’, ‘partly-compliant’ or ‘non-compliant’. Horses exhibited a decreased HR (P = 0.015) and a decreased BS (P = 0.004) within and across different tests. ‘Matching’ horse-rider combinations exhibited less stress as indicated by reduced HR (‘match’ 69 ± 10 vs. ‘mismatch’ 72 ± 9, P = 0.001) and BS (‘match’ 1.9 ± 1.1 vs. ‘mismatch’ 3.8 ± 1.4, P = 0.017) of the horse. ‘Compliant’ (68 ± 8, P < 0.001) and ‘partly-compliant’ (71 ± 9, P = 0.002) horses had significantly lower HR than ‘non-compliant’ (75 ± 9) animals. The findings of the study indicate that HR and BS measurements support a subjective ‘match’ diagnosis and HR measurement may be a valuable tool in assessing horse compliance.
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König v. Borstel, U., Visser, E. K., & Hall, C. (2017). Indicators of stress in equitation. Appl. Anim. Behav. Sci., 190, 43–56.
Abstract: Abstract Stress is a generic concept describing the body's reaction to external stimuli, including both physiological and psychological factors. Therefore, by definition, the assessment of psychological stress in the exercising horse encompasses the problem of teasing apart the psychological and physiological factors both of which result in stress responses. The present study reviews the existing literature on various measures of stress taken specifically in the context of equitation science. Particular attention has been paid to short-term effects, and commonly used measurements of short-term stress include heart rate, a number of heart rate variability parameters, blood or saliva cortisol levels, eye temperature, and various behaviour parameters including in particular behaviour patterns presumably indicative of conflict with the rider's/trainer's aids. Inspection of the individual studies' results revealed that disagreement between these different measures of stress is commonplace. For physiological parameters, the largest proportion of agreement (i.e. both parameters simultaneously indicated either higher, insignificant or lower stress compared to a control treatment) was found for heart rate and heart rate variability parameters, while generally limited agreement was found for cortisol. It appears that cortisol levels may not be particularly useful for assessing/assessment of the valence of a situation in the exercising horse as cortisol levels are predominantly linked to activation and exercise levels. Although heart rate variability parameters reflect in theory more closely sympathovagal balance compared to cortisol levels, great care has to be taken regarding the use of appropriate time-frames, appropriate raw data correction methods as well as the use of appropriate equipment. In spite of its wide-spread and apparently successful use, popular equipment may in fact not be accurate enough under field conditions. Eye temperature is another promising parameter for assessment of psychological stress, but the technique is likewise susceptible to application errors. Given the high susceptibility of physiological parameters to errors at various experimental stages, behavioural rather than physiological parameters may in fact provide more accurate measures of valence when conducting experiments in the exercising horse. Behavioural parameters that appear to be particularly practical in assessing stress in ridden horses' behaviour are associated with frequencies of behaviour indicative of conflict. However, while increased frequencies of are a good indicator of stress, the absence of conflict behaviour does not provide proof of the absence of stress due to the possible occurrence of conditions such as Learned Helplessness. In future studies, the above issues should be taken into consideration when designing experiments to assess psychological stress in ridden horses.
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McLean, A. N., & Christensen, J. W. (2017). The application of learning theory in horse training. Appl. Anim. Behav. Sci., 190, 18–27.
Abstract: The millennia-old practices of horse training markedly predate and thus were isolated from the mid-twentieth century revelation of animal learning processes. From this standpoint, the progress made in the application and understanding of learning theory in horse training is reviewed including a discussion of how learning processes are employed or otherwise under-utilised in training. This review describes the process of habituation and the most commonly applied desensitisation techniques (systematic desensitisation, counter-conditioning, overshadowing, response prevention) and propose two additional techniques (approach conditioning and stimulus blending). The salience of different types of cues, the interaction of operant and classical conditioning and the impact of stress are also discussed. This paper also exposes the inflexibility and occasional inadequacy of the terminology of learning theory when translated from the research laboratory situation to the practical setting in horse training. While learning theory provides a rich toolbox for riders and trainers, the training process is subject to the simultaneous use of multiple learning processes. In addition, learning/behavioural outcomes and trained responses are not just the result of simple stimulus-response based interactions but are further shaped by arousal, affective and attachment states. More research is needed in these areas. For the field of equitation science to progress and to improve clarity and use of learning processes, changes in nomenclature are required. In particular, the use of the terms 'positive' and 'negative' as descriptive labels in both reinforcement and punishment modalities are unacceptably misleading for everyday use. These labels inhibit the understanding and recognition of the learning processes that these terms supposedly represent, yet the learning processes they describe are vital for horse riders, handlers and trainers to understand. We therefore propose that these labels should be re-labelled more appropriately as 'addition' or 'subtraction' reinforcement/punishment. This would enlighten trainers on the correct application of learning theory, and safety and welfare benefits for people and horses would follow. Finally it is also proposed that the term 'conflict theory' be taken up in equitation science to facilitate diagnosis of training-related behaviour disorders and thus enable the emergence of improved training practices. The optimal use of learning theory should be established as a fundamental principle in equestrian education.
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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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