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Nagy, K., Bodó, G., Bárdos, G., Harnos, A., & Kabai, P. (2009). The effect of a feeding stress-test on the behaviour and heart rate variability of control and crib-biting horses (with or without inhibition). Appl. Anim. Behav. Sci., 121(2), 140–147.
Abstract: Crib-biting is a form of oral stereotypy affecting 4-5% of horses. Once fixed, crib-biting is difficult to eliminate by behaviour therapy, however, its performance can be inhibited by collar or surgery treatment (modified Forssell's procedure). Although surgical intervention is widespread, the effects on stress coping in horses have not been studied. In the present study we evaluated changes in behaviour response and heart rate variability in 9 control, 10 crib-biting, 10 collar and 11 surgically treated horses in a feeding stress-test, in which a feeding-bowl was placed in front but out of the reach of the horses, from which tidbits were given 3 times. We found that stress triggers high oral activity, mainly cribbing in crib-biting horses, elevates other forms of oral activities in the inhibited groups and does not affect oral activities of controls. Instead of performing oral activities, control horses tended to target an unavailable feeding-bowl by pawing or head-tossing. Changes in stress level were indistinguishable in controls and crib-biters as heart rate variability returned to baseline values in both groups. In contrast, horses inhibited to perform crib-biting showed elevated stress level throughout the test period. Our results suggest that crib-biting may develop to cope with stress, and such coping function diminishes when inhibited.
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Kay, R., & Hall, C. (2009). The use of a mirror reduces isolation stress in horses being transported by trailer. Appl. Anim. Behav. Sci., 116(2-4), 237–243.
Abstract: Horse trailers are a common form of transportation for horses and ponies and often require the animal to travel alone or with a single companion. The current study investigated the effect of transporting horses alone, in company or with an acrylic safety mirror (measuring 81cm61.5cm) that provided surrogate companionship. The behavioural and physiological responses of 12 mature horses during a 30-min journey by trailer under the three treatments were compared. Behaviours (vocalisation, eating, head-tossing, pawing, and head-turning) were recorded. In order to assess circulatory changes that occur as part of the response to transport, heart rate (HR), rectal (Tr) and ear-pinna (Tp) temperatures were recorded. When travelling with a live companion significantly less time was spent vocalising (p<0.001), head-turning (p<0.001), head-tossing (p<0.01) and pawing (p<0.01); eating behaviour increased (p<0.05). Physiological responses (increases in HR and Tr and decreases in Tp) were also significantly reduced when travelling with a live companion (p<0.01). Travelling with the mirror did not significantly affect physiological responses compared with travelling alone, but the rise in Tr and fall in Tp was reduced (p=0.052 and p=0.051, respectively) and can be considered a trend. When travelling with a mirror significantly less time was spent turning the head (p<0.01), vocalising (p<0.05) and head-tossing (p<0.05); eating behaviour increased (p<0.05). The only significant difference between travelling with a live companion and a mirror was that the time spent turning the head round was less with a live companion (p<0.05). The provision of surrogate companionship in the form of a mirror was found to be preferable to travel alone, but where possible a live companion is recommended. Isolation during transportation was found to suppress feeding behaviour. Although peripheral blood flow (Tp) has been used to assess transport stress in other species it has not previously been used in the horse. Further evaluation of this non-invasive measure is now required.
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Rogers, L. J. (2010). Relevance of brain and behavioural lateralization to animal welfare. Appl. Anim. Behav. Sci., 127(1-2), 1–11.
Abstract: The left and right sides of the brain are specialised to process information in different ways and to control different categories of behaviour. Research on a range of species has shown that the left hemisphere controls well-established patterns of behaviour performed in non-stressful situations, whereas the right hemisphere responds to unexpected stimuli and controls escape and other emergency responses. The known functions of each hemisphere are summarised in this paper. Then it is hypothesised that stressed animals rely on predominant use of the right hemisphere, and that a bias to use the right or left hemisphere, respectively, may explain the behavioural differences between animals with a negative cognitive bias and those with a positive cognitive bias. In some species of primates it has been shown that the preferred limb used to pick up food when the animal is in a relaxed state reflects the dominant hemisphere and may be an accessible measure indicating susceptibility to stress and tendency towards positive versus negative cognitive bias. Hence, limb preference might be a useful measure of such tendencies in domesticated species. Some difficulties in determining a relevant measure of limb preference in non-primate species are mentioned, followed by the suggestion that eye preferences for viewing certain stimuli may be a useful measure in species with laterally placed eyes. Finally, effects of experience on the development of hemispheric dominance are discussed, leading to a suggestion that the welfare of domestic animals may be enhanced by ensuring development of left hemisphere dominance (e.g. by exposing chick embryos to light) and by shifting right to left hemisphere dominance in animals with negative cognitive bias.
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Innes, L., & McBride, S. (2008). Negative versus positive reinforcement: An evaluation of training strategies for rehabilitated horses. Appl. Anim. Behav. Sci., 112(3–4), 357–368.
Abstract: Rescued equids are often exposed to rehabilitation and training (or retraining) programmes to improve their physical and psychological well-being as well as to facilitate the re-homing process. Training uses either positive or negative reinforcement learning procedures and it is considered here that, there may be welfare implications associated with using the latter technique as it has the potential to overlay acute stress on animals with a chronic stress life history. The aim of this study, therefore, was to compare these training strategies (negative versus positive reinforcement) on equine behaviour and physiology as the first step in establishing an optimal rehabilitation approach (from a welfare perspective) for equids that have been subjected to chronic stress in the form of long-term neglect/cruelty. Over a 7-week period, 16 ponies (aged 6–18 months) were trained using either positive (‘positive’) (n = 8) or negative reinforcement (‘negative’) (n = 8) techniques to lead in hand, stand to be groomed, traverse an obstacle course and load into a trailer. Heart rate was measured (5 s intervals) on days 1 and 4 of each training week, ‘Pre’- (1 h), ‘During’ (0.5 h) and ‘Post’- (1 h) training session. Ethograms (10.00–20.00 h) outside of the training period were also compiled twice weekly. In addition, weekly arena tests (as a measure of reactivity) were also performed 1 week before and during the 7 weeks of training. Results showed significant differences between the two training schedules for some measures during the latter stages of the trial and suggested that animals trained under a positive reinforcement schedule were more motivated to participate in the training sessions and exhibited more exploratory or ‘trial and error’ type behaviours in novel situations/environments. In this context, the incorporation of positive reinforcement schedules within a rehabilitation programme may be of benefit to the animal from a welfare perspective.
Keywords: Horse; Training; Positive reinforcement; Negative reinforcement; Stress; Rehabilitation
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Christensen, J. W., Ahrendt, L. P., Lintrup, R., Gaillard, C., Palme, R., & Malmkvist, J. (2012). Does learning performance in horses relate to fearfulness, baseline stress hormone, and social rank? In Applied Animal Behaviour Science (Vol. 140, pp. 44–52).
Abstract: The ability of horses to learn and remember new tasks is fundamentally important for their use by humans. Fearfulness may, however, interfere with learning, because stimuli in the environment can overshadow signals from the rider or handler. In addition, prolonged high levels of stress hormones can affect neurons within the hippocampus; a brain region central to learning and memory. In a series of experiments, we aimed to investigate the link between performance in two learning tests, the baseline level of stress hormones, measured as faecal cortisol metabolites (FCM), fearfulness, and social rank. Twenty-five geldings (2 or 3 years old) pastured in one group were included in the study. The learning tests were performed by professional trainers and included a number of predefined stages during which the horses were gradually trained to perform exercises, using either negative (NR) or positive reinforcement (PR). Each of the learning tests lasted 3 days; 7min/horse/day. The NR test was repeated in a novel environment. Performance, measured as final stage in the training programme, and heart rate (HR) were recorded. Faeces were collected on four separate days where the horses had been undisturbed at pasture for 48h. Social rank was determined through observations of social interactions during feeding. The fear test was a novel object test during which behaviour and HR were recorded. Performance in the NR and PR learning tests did not correlate. In the NR test, there was a significant, negative correlation between performance and HR in the novel environment (rS=-0.66, P<0.001, i.e. nervous horses had reduced performance), whereas there was no such correlation in the home environment (both NR and PR). Behavioural reactions in the fear test correlated significantly with performance in the NR test in the novel environment (e.g. object alertness and final stage: rS=-0.43, P=0.04), suggesting that performance under unfamiliar, stressful conditions may be predicted by behavioural responses in a fear test. There was a negative correlation between social rank and baseline stress hormones (rS=-0.43, P=0.04), i.e. high rank corresponded to low FCM concentrations, whereas neither rank nor FCM correlated with fearfulness or learning performance. We conclude that performance under stressful conditions is affected by activation of the sympathetic nervous system during training and related to behavioural responses in a standardised fear test. Learning performance in the home environment, however, appears unrelated to fearfulness, social rank and baseline FCM levels.
Keywords: Horse; Learning; Fearfulness; Stress; Reinforcement; Social rank
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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.
Keywords: Behaviour; Conflict theory; Desensitisation; Habituation; Reinforcement; Stress
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Davies, H. M. S. (2005). The timing and distribution of strains around the surface of the midshaft of the third metacarpal bone during treadmill exercise in one Thoroughbred racehorse. Aust Vet J, 83(3), 157–162.
Abstract: OBJECTIVE: To confirm that the midshaft dorsal cortex of the third metacarpal bone experienced higher compressive strains during fast exercise than the medial or lateral cortices, and that the strain peak occurred earlier in the hoof-down phase of the stride on the dorsal cortex than the medial or lateral cortices. DESIGN: Observations of a single horse. PROCEDURE: Strains were collected from a single, sound, 3-year-old Thoroughbred mare during treadmill exercise from rosette strain gauges implanted onto the medial, lateral and dorsal surfaces of the midshaft of the right cannon bone, simultaneously with data from a hoof switch that showed when the hoof was in the stance phase. RESULTS: Peak compressive strains on the dorsal surface of the third metacarpal bone were proportional to exercise speed and occurred at about 30% of stance. Peak compressive strains on the medial surface of the non-lead limb reached a maximum at a speed around 10 m/s and occurred at mid-stance. Peak compressive strains on the lateral surface varied in timing and size between strides at all exercise speeds, but remained less than -2000 microstrains. CONCLUSIONS: The timing of peak compressive strains on the dorsal cortex suggests a relationship to deceleration of the limb following hoof impact, so the main determinants of their size would be exercise speed and turning (as shown in previous experiments). This experiment confirms data from other laboratories that were published but not discussed, that peak compressive strains on the medial surface occur at mid-stance. This suggests that they are related to the support of body weight. The strains on the lateral cortex occurred at variable times so may be associated with the maintenance of balance as well as the support of body weight. Understanding the loading of the third metacarpal bone will help to determine causes of damage to it and ways in which the bone might be conditioned to prevent such damage.
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Massen, J., Sterck, E., & de Vos, H. (2010). Close social associations in animals and humans: functions and mechanisms of friendship (Vol. 147).
Abstract: Both humans and group-living animals associate and behave affiliatively more with some individuals than others. Human friendship has long been acknowledged, and recently scientists studying animal behaviour have started using the term friendship for close social associates in animals. Yet, while biologists describe friends as social tools to enhance fitness, social scientists describe human friendship as unconditional. We investigate whether these different descriptions reflect true differences in human friendship and animal close social associations or are a by-product of different research approaches: namely social scientists focussing on proximate and biologists on ultimate explanations. We first stress the importance of similar measures to determine close social associations, thereafter examine their ultimate benefits and proximate motivations, and discuss the latest findings on the central-neural regulation of social bonds. We conclude that both human friendship and animal close social associations are ultimately beneficial. On the proximate level, motivations for friendship in humans and for close social associations in animals are not necessarily based on benefits and are often unconditional. Moreover, humans share with many animals a similar physiological basis of sociality. Therefore, biologists and social scientist describe the same phenomenon, and the use of the term friendship for animals seems justified.
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Hoshaw, B. A., Evans, J. C., Mueller, B., Valentino, R. J., & Lucki, I. (2006). Social competition in rats: Cell proliferation and behavior. Behav. Brain. Res., 175(2), 343–351.
Abstract: Behavioral and physiological changes were studied following prolonged exposure to social competition in pairs of non-food-deprived rats competing daily for a limited supply of graham cracker crumbs. Stable dominant-subordinate relationships developed in most pairs, as measured by feeding time, which were maintained over a 5-6-week study period. In other behavioral tests, subordinates demonstrated a decreased latency to immobility in the forced swim test compared with dominants, but no difference in locomotor activity. Subordinates had increased bladder size, decreased adrenal gland size, and a 35% reduction of hippocampus cell proliferation compared with the dominant member. Therefore, prolonged social competition, based on restricted access to palatable substances, produced hierarchies among individuals that were associated with differences in behavior, physiology and hippocampal cell proliferation.
Keywords: Social stress; Depression; Forced swim test; Neurogenesis
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