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Nuñez, C. M. V., Adelman, J. S., Smith, J., Gesquiere, L. R., & Rubenstein, D. I. (2014). Linking social environment and stress physiology in feral mares (Equus caballus): Group transfers elevate fecal cortisol levels. General and Comparative Endocrinology, 196, 26–33.
Abstract: Abstract Feral horses (Equus caballus) have a complex social structure, the stability of which is important to their overall health. Behavioral and demographic research has shown that decreases in group (or band) stability reduce female fitness, but the potential effects on the physiological stress response have not been demonstrated. To fully understand how band stability affects group-member fitness, we need to understand not only behavioral and demographic, but also physiological consequences of decreases to that stability. We studied group changes in feral mares (an activity that induces instability, including both male and female aggression) on Shackleford Banks, NC. We found that mares in the midst of changing groups exhibit increased fecal cortisol levels. In addition, mares making more group transfers show higher levels of cortisol two weeks post-behavior. These results offer insights into how social instability is integrated into an animal’s physiological phenotype. In addition, our results have important implications for feral horse management. On Shackleford Banks, mares contracepted with porcine zona pellucida (PZP) make approximately 10 times as many group changes as do untreated mares. Such animals may therefore be at higher risk of chronic stress. These results support the growing consensus that links between behavior and physiological stress must be taken into account when managing for healthy, functional populations.
Keywords: Equus caballus; Fecal cortisol; Feral mare; Group transfer; Stress; Social instability
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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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Nagy, K., Bodó, G., Bárdos, G., & Harnos, A. (2008). Is modified Forssell"s operation superior to cribbing collar in preventing crib-biting in horses? In IESM 2008.
Abstract: Crib-biting (wind-sucking) might be a coping response of the horses to the challenges of
uncontrolled environmental events. Prevention of this stereotypic behaviour evokes physiological responses consistent with increased stress. Reducing the incidence of cribbiting, however, is important in order to prevent undesirable physical and behavioural consequences (tooth erosion, altered gut function, gastric inflammation/ulceration, colic, etc.). Common treatment of crib-biting is the application of a cribbing collar, which limits the flexion of the neck making this stereotypic movement uncomfortable and difficult. Another method, the modified Forssell"s operation, is becoming more and more popular amongst the horse owners. It is based on the removal of the muscles used in crib-biting (m.omohyoideus, m.sternohyoideus, m.sternothyrohyoideus) and the ventral branches of the spinal accessory nerves. Surveys on the success of this surgical procedure have revealed inconsistent results, and, contrary to the cribbing collar, its effect on the stress level have not been studied either. The aim of our study was to determine whether the modified Forssell"s procedure is superior to the cribbing collar treatment. Differences in stress management was tested by a crib-biting provoking test, in which surgically treated horses, crib-biting horses, crib-biting horses with cribbing collar, and normal horses (those showing no stereotypies), altogether 56 horses were compared. In this test, a food bucket had been placed out of the reach of the animal, from which titbits were given 3 times. Behaviour and heart rate variability (HRV) of the horses were recorded and analysed throughout the test. Hypotheses were tested by linear mixed model. According to our results, both prevention methods (collar or surgery) inhibited crib-biting successfully though not totally. Regarding behaviour and heart rate variability, horses prevented from crib-biting (by collar or surgery) differed significantly from crib-biting and normal horses but not from each other. Normal horses were usually trying to reach the food-bucket while present and were standing still afterwards, whereas the other three groups had not really made efforts to reach the bucket, spent less time with resting, and performed or tried crib-biting. During the stress-test, normal and crib-biting horses had shown good stress-adaptation to the challenge since their HRV, after an initial increase, returned to the basal value by the end. On the contrary, HRV of the two prevented groups remained elevated and showed large oscillations throughout. They had not found a successful coping behaviour either. Our results suggest that since prevention may significantly increase distress, the treatment in itself, without changing the motivation of the horse to perform the replacement behaviour – it seems to be unsatisfactory and insufficient. After prevention the motivation of the horse to perform crib-biting should be addressed. In addition, considering that prevention by collar and surgery had not resulted in any significant behavioural or physiological differences, the superiority of the modified Forssell"s procedure might be questioned. However, the surgery might be recommended if treatment with collar is ineffective. |
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Mormède, P., Andanson, S., Aupérin, B., Beerda, B., Guémené, D., Malmkvist, J., et al. (2007). Exploration of the hypothalamic-pituitary-adrenal function as a tool to evaluate animal welfare. Physiol. Behav., 92(3), 317–339.
Abstract: Measuring HPA axis activity is the standard approach to the study of stress and welfare in farm animals. Although the reference technique is the use of blood plasma to measure glucocorticoid hormones (cortisol or corticosterone), several alternative methods such as the measurement of corticosteroids in saliva, urine or faeces have been developed to overcome the stress induced by blood sampling itself. In chronic stress situations, as is frequently the case in studies about farm animal welfare, hormonal secretions are usually unchanged but dynamic testing allows the demonstration of functional changes at several levels of the system, including the sensitization of the adrenal cortex to ACTH and the resistance of the axis to feedback inhibition by corticosteroids (dexamethasone suppression test). Beyond these procedural aspects, the main pitfall in the use of HPA axis activity is in the interpretation of experimental data. The large variability of the system has to be taken into consideration, since corticosteroid hormone secretion is usually pulsatile, follows diurnal and seasonal rhythms, is influenced by feed intake and environmental factors such as temperature and humidity, age and physiological state, just to cite the main sources of variation. The corresponding changes reflect the important role of glucocorticoid hormones in a number of basic physiological processes such as energy metabolism and central nervous system functioning. Furthermore, large differences have been found across species, breeds and individuals, which reflect the contribution of genetic factors and environmental influences, especially during development, in HPA axis functioning. Usually, these results will be integrated with data from behavioral observation, production and pathology records in a comprehensive approach of farm animal welfare.
Keywords: Stress; Animal welfare; HPA axis; Glucocorticoid hormones; Acth; Dexamethasone suppression test; Cattle; Pig; Fur animals; Mink; Fox; Poultry; Fish
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Minero, M., Canali, E., Ferrante, V., Verga, M., & Odberg, F. O. (1999). Heart rate and behavioural responses of crib-biting horses to two acute stressors. Vet. Rec., 145(15), 430–433.
Abstract: The heart rate and behaviour of 14 adult saddle horses, eight crib-biters and six normal controls, were investigated. Initially, the relationship between crib-biting and heart rate was investigated while the horses were undisturbed. The horses were tested when restrained with a lip twitch, and assessed when they were exposed suddenly to the rapid inflation of a balloon. The heart rate of the crib-biters during crib-biting was lower than during other behaviours. The crib-biters had a higher overall mean heart rate (P<0.05) suggesting that they may have had a higher basal sympathetic activity. After the application of the twitch, all the horses had a transient increase in heart rate which returned to basal values more rapidly in the crib-biters. The crib-biters were less reactive to the lip twitch, five of the six investigated remaining calm, and after the release of the twitch, they spent more time nibbling (P<0.05) than the control horses. The crib-biters reacted more strongly to the inflation of the balloon (three of the six reacted), and after it had been inflated they spent more time walking in the box.
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Mills, D. S., Alston, R. D., Rogers, V., & Longford, N. T. (2002). Factors associated with the prevalence of stereotypic behaviour amongst Thoroughbred horses passing through auctioneer sales. Appl. Anim. Behav. Sci., 78(2-4), 115–124.
Abstract: The objective of this study was to evaluate whether sex, age and/or coat colour were associated with the occurrence of stereotypic behaviour in the horse and to assess whether the occurrence of one type of stereotypy in an individual was associated with the occurrence of another specific type of stereotypy. The incidence of stereotypic boxwalking, weaving (both locomotor stereotypies) and oral stereotypy in 4061 Thoroughbred horses passing through five bloodstock auctions were recorded from sale declarations and information on returns. An overall prevalence of 5.1% was recorded, and varied with sex (P<0.001) and age (P<0.001) but not coat colour (P=0.495). Prevalence was higher in females, geldings, and 2-year-olds. Examination of the assumption that stereotypies are acquired independently suggested a higher than expected prevalence of animals with more than one stereotypy. The interaction was not the same for all forms of stereotypy recorded. The effect was greatest between boxwalking and weaving, (odds ratio 13.6) whilst combinations involving oral and locomotor stereotypies had lower odds ratios (between 2.9 and 4.9).
Keywords: Behaviour; Horses; Management; Prevalence; Stereotypy; Stress
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Mendl, M. (1999). Performing under pressure: stress and cognitive function. Appl. Anim. Behav. Sci., 65(3), 221–244.
Abstract: The way in which cognitive functioning is affected by stressors is an important area of research for applied ethologists because stress caused by captive conditions may disrupt cognitive processes and lead to welfare and husbandry problems. Such problems may be minimised through an understanding of the links between stress and cognition. The effects of stress on cognitive function have been studied in disciplines ranging from human perceptual psychology to animal neuroscience. The aim of this paper is to provide an introduction to this research, focusing on the effects of stressors on attention, memory formation and memory recall. Findings from such a diverse literature with little apparent inter-disciplinary communication are inevitably complex and often contradictory. Nevertheless, some generalities do emerge. The idea that an inverted U-shaped relationship exists between an individual's state of stress or arousal and its ability to perform a cognitive task effectively, the so-called Yerkes-Dodson law, is commonly encountered. The law has limited explanatory value because it is unlikely that different stressors act on cognitive function via the same intervening, non-specific state. Furthermore, the law only provides a very general description of the relationship between stress and cognitive function. Empirical research on attention and memory processes reveals more specific findings. Stressors appear to cause shifts, lapses and narrowing of attention, and can also influence decision speed. These processes may be viewed as serving an adaptive role helping the animal to search for and scrutinise a source of danger. There is conflicting evidence as to whether hormones involved in the hypothalamic-pituitary-adrenal stress response play a part in these processes. These hormones and those involved in the sympathetic-adrenomedullary stress response do appear to play an important role in memory formation. Low or moderate concentrations of circulating glucocorticoids and catecholamines can enhance memory formation, while excessively high or prolonged elevations of these hormones can lead to memory disruption. The effects of stressors on memory recall are less clear. There is evidence for disruptive effects, and for facilitatory effects indicating state-dependent memory recall; events experienced under conditions of high arousal may be best recalled under similar conditions. Applied ethologists have the opportunity to extend work in this area, which often involves studies of single stressors/stress hormones acting in isolation and limited measures of cognitive function, by focusing on real-life husbandry stressors encountered by captive animals. This will yield fundamental information which also has direct relevance to animal welfare and management issues.
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Meehan, C. L., & Mench, J. A. (2007). The challenge of challenge: Can problem solving opportunities enhance animal welfare? Appl. Anim. Behav. Sci., 102(3-4), 246–261.
Abstract: Cognitive mechanisms are an important part of the organization of the behavior systems of animals. In the wild, animals regularly face problems that they must overcome in order to survive and thrive. Solving such problems often requires animals to process, store, retrieve, and act upon information from the environment--in other words, to use their cognitive skills. For example, animals may have to use navigational, tool-making or cooperative social skills in order to procure their food. However, many enrichment programs for captive animals do not include the integration of these types of cognitive challenges. Thus, foraging enrichments typically are designed to facilitate the physical expression of feeding behaviors such as food-searching and food consumption, but not to facilitate complex problem solving behaviors related to food acquisition. Challenging animals by presenting them with problems is almost certainly a source of frustration and stress. However, we suggest here that this is an important, and even necessary, feature of an enrichment program, as long as animals also possess the skills and resources to effectively solve the problems with which they are presented. We discuss this with reference to theories about the emotional consequences of coping with challenge, the association between lack of challenge and the development of abnormal behavior, and the benefits of stress (arousal) in facilitating learning and memory of relevant skills. Much remains to be done to provide empirical support for these theories. However, they do point the way to a practical approach to improving animal welfare--to design enrichments to facilitate the cognitive mechanisms which underlie the performance of complex behaviors that cannot be performed due to the restrictions inherent to the captive environment.
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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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McGee, S., & Smith, H. V. (2004). Accompanying pre-weaned Thoroughbred (Equus caballus) foals while separated from the mare during covering reduces behavioural signs of distress exhibited. Appl. Anim. Behav. Sci., 88(1-2), 137–147.
Abstract: Some Thoroughbred stallion farms separate the mare from the foal during breeding, leaving the foal alone in its stall. Such separation often leads to distress in the foal, as evidenced by behavioural indicators such as vocalisations, pacing or moving about, pawing and striking the body against solid objects. This study examined whether the foal's distress is reduced if a handler accompanies it during the period of separation. A total of 57 foals on an Irish stud farm were observed: 27 foals were accompanied during separation and 30 were left in isolation but discretely observed from outside the stall. On average, unaccompanied foals vocalised at significantly (P<0.001) shorter intervals (14.8 s versus 26.8 s), and spent significantly (P<0.05) more time pawing (29.6 s versus 6.8 s) than accompanied ones. In addition, 17 of the unaccompanied foals hurled themselves against a solid object on at least one occasion, while not one of the accompanied foals did this (P<0.001). Age, gender and number of times the foal had been previously separated from the mare while she was being teased or covered were not significantly related to any of the behavioural indicators of distress. Overall, the data provide clear evidence that accompanying a foal reduces the distress that it exhibits when separated from the mare. This procedure has obvious welfare benefits in the short term, in that the foal both experiences less distress and has a reduced likelihood of injuring itself, and it may have longer-term benefits by reducing the chance of the animal later developing a locomotory stereotypical behaviour.
Keywords: Distress; Foals; Maternal separation; Stereotypies; Welfare
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