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VanDierendonck, M. C. (2006). General Discussion (Vol. Chapter 7). Universiteit Utrecht.
Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
Keywords: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
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Bachmann, I., Bernasconi, P., Herrmann, R., Weishaupt, M. A., & Stauffacher, M. (2003). Behavioural and physiological responses to an acute stressor in crib-biting and control horses. Appl. Anim. Behav. Sci., 82(4), 297–311.
Abstract: The responses of eleven pairs of crib-biting and non-crib-biting horses (controls) to an arousal-inducing stimulus were studied. Video-observation of the horses revealed that crib-biting horses spent between 10.4 and 64.7% of their stabling time performing the stereotypy. During the first 2 days of an experimental period, the horses were conditioned to receive food from a special bucket. On the third day the food bucket was presented, but the horses were not allowed to feed. Arousal behaviour and crib-biting intensity as well as plasma cortisol concentration, heart rate (HR) and heart rate variability (HRV) were recorded at rest, and during and after presentation of the food stimulus. The stimulus induced a significant increase of HR and arousal behaviour in crib-biters and in controls, whereas the crib-biting frequency decreased. Power spectral analysis of the HRV revealed significant differences between crib-biters and controls at rest: crib-biters had a lower vagal tone (high frequency component, HF) and a higher sympathetic tone (low frequency component, LF) than controls. The lower basal parasympathetic activity might be an indication why crib-biting horses, in contrast to the controls, showed neither a significant decrease of the HF component during presentation of the food stimulus nor an increase of the HF component after presentation. Thus, there might be differences in the tuning of the autonomous nervous system and of the stress reactivity in crib-biting and in control horses. The results suggest that the crib-biting horses are more stress sensitive and physiologically and psychologically less flexible than the control horses.
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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).
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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.
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Art, T., & Lekeux, P. (2005). Exercise-induced physiological adjustments to stressful conditions in sports horses. Adaptability of sport horses to stressful conditions, 92(2), 101–111.
Abstract: Among athletic/sports animals, the horse has a unique ability to increase its oxygen uptake by a factor of 60 during heavy exercise. This is achieved by physiological adaptations of all the links in the oxygen chain. Ventilation is increased by a factor of 30. Since the horse is a compulsory nasal breather, this hyperpnea necessitates high transmural pressure changes, which may be responsible for the dynamic collapse of the airways. Blood flow is increased by a factor of 10. Since the left ventricle is not very compliant, this increase necessitates a high filling pressure in the pulmonary circulation, which may induce capillary stress failure and exercise-induced pulmonary haemorrhage. Lastly, oxygen transport is improved by splenic contraction which increases haemoglobinemia by 50%. Sports horses frequently suffer from several problems, which are related either to endogenous or exogenous stresses experienced during their career. These stresses, caused by the use of the horse as a competition animal, may lead to several medical problems. At a systemic level, endogenous stresses include hyperkaliemia, lactacidemia, and hyperthermia; oxidative stress may induce problems at a general, and/or a pulmonary level. External factors, e.g. poor quality of inspired air, transport, hot and humid ambient conditions, and microbiological agents, may also induce abnormal body attacks, and lead to health problems.
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Oikawa, M., Hobo, S., Oyamada, T., & Yoshikawa, H. (2005). Effects of Orientation, Intermittent Rest and Vehicle Cleaning During Transport on Development of Transport-related Respiratory Disease in Horses. Journal of Comparative Pathology, 132(2-3), 153–168.
Abstract: Summary The effects of various factors on the inflammatory and stress response in horses during transportation were examined in Experiments 1 and 2, carried out in April and August, respectively. In Experiment 1, three groups (G1-G3) of four Thoroughbreds were used, and in Experiment 2, two groups (G4, G5). G1 animals were loaded into lorries with their heads facing forwards (FF) and given periods of short rest (SR) (30 min for every 4 h driven). G2 horses were loaded facing rearwards (FR) and given SR. G3 horses were FF and given periods of long rest (LR) (2 h rest for every 4 h driven). G4 horses were transported FF with hay suspended in front of them and no cleaning of faeces or urine during rest stops. G5 horses were FF and given pellets instead of hay, and the lorry was cleaned and washed at each rest stop. G4 and G5 horses were rested for 1 h after every 5 h of driving. All groups were driven a distance of 1500 km, and the total journey time was 37 h for G1 and G2, 49 h for G3, and 40 h for G4 and G5. In Experiment 1, indicators of inflammatory and stress responses tended to be highest in G1, intermediate in G2 and lowest in G3. In Experiment 2 they tended to be higher in G4 than in G5. The results suggested that increasing the rest time and cleaning the interior of the vehicle during rest stops reduced transportation stress and respiratory insults, factors that may lead to respiratory disease.
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Stahl, F., & Dorner, G. (1982). Responses of salivary cortisol levels to stress-situations. Endokrinologie, 80(2), 158–162.
Abstract: A procedure is described for determining salivary cortisol levels by a competitive protein-binding assay using horse transcortin. The collection of saliva was performed by means of filter paper-strips. Filter paper samples are more than 5 days stable after air-drying. In this form, the samples could be stored without refrigerator or deep-freezer and, if necessary, sent by post to the laboratory without any special precaution. Stressful situation of either painful or anxious origin were associated with an adequate increase of salivary cortisol levels. The increases were 157 to 230% of the initial or normal values dependent on the kind of stress. The mean values in 4 cases of Cushing's syndrome were 380% and 1 hour after 25 I.U. ACTH 690% higher than those in normal persons. In normal persons, a well-defined circadian rhythm has been observed.
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Creel, S. (2001). Social dominance and stress hormones. Trends. Ecol. Evol, 16(9), 491–497.
Abstract: In most cooperatively breeding birds and mammals, reproductive rates are lower for social subordinates than for dominants, and it is common for reproduction in subordinates to be completely suppressed. Early research conducted in captivity showed that losing fights can increase glucocorticoid (GC) secretion, a general response to stress. Because GCs can suppress reproduction, it has been widely argued that chronic stress might underlie reproductive suppression of social subordinates in cooperative breeders. Contradicting this hypothesis, recent studies of cooperative breeders in the wild show that dominant individuals have elevated GCs more often than do subordinates. The findings that elevated GCs can be a consequence of subordination or a cost of dominance complicate the conventional view of social stress, with broad ramifications for the evolution of dominance and reproductive suppression.
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Palme, R., Rettenbacher, S., Touma, C., El-Bahr, S. M., & Mostl, E. (2005). Stress hormones in mammals and birds: comparative aspects regarding metabolism, excretion, and noninvasive measurement in fecal samples. Ann N Y Acad Sci, 1040, 162–171.
Abstract: A multitude of endocrine mechanisms are involved in coping with challenges. Front-line hormones to overcome stressful situations are glucocorticoids (GCs) and catecholamines (CAs). These hormones are usually determined in plasma samples as parameters of adrenal activity and thus of disturbance. GCs (and CAs) are extensively metabolized and excreted afterwards. Therefore, the concentration of GCs (or their metabolites) can be measured in various body fluids or excreta. Above all, fecal samples offer the advantages of easy collection and a feedback-free sampling procedure. However, large differences exist among species regarding the route and time course of excretion, as well as the types of metabolites formed. Based on information gained from radiometabolism studies (reviewed in this paper), we recently developed and successfully validated different enzyme immunoassays that enable the noninvasive measurement of groups of cortisol or corticosterone metabolites in animal feces. The determination of these metabolites in fecal samples can be used as a powerful tool to monitor GC production in various species of domestic, wildlife, and laboratory animals.
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Touma, C., Sachser, N., Mostl, E., & Palme, R. (2003). Effects of sex and time of day on metabolism and excretion of corticosterone in urine and feces of mice. Gen Comp Endocrinol, 130(3), 267–278.
Abstract: Non-invasive techniques to monitor stress hormones in small animals like mice offer several advantages and are highly demanded in laboratory as well as in field research. Since knowledge about the species-specific metabolism and excretion of glucocorticoids is essential to develop such a technique, we conducted radiometabolism studies in mice (Mus musculus f. domesticus, strain C57BL/6J). Each mouse was injected intraperitoneally with 740 kBq of 3H-labelled corticosterone and all voided urine and fecal samples were collected for five days. In a first experiment 16 animals (eight of each sex) received the injection at 9 a.m., while eight mice (four of each sex) were injected at 9 p.m. in a second experiment. In both experiments radioactive metabolites were recovered predominantly in the feces, although males excreted significantly higher proportions via the feces (about 73%) than females (about 53%). Peak radioactivity in the urine was detected within about 2h after injection, while in the feces peak concentrations were observed later (depending on the time of injection: about 10h postinjection in experiment 1 and about 4h postinjection in experiment 2, thus proving an effect of the time of day). The number and relative abundance of fecal [3H]corticosterone metabolites was determined by high performance liquid chromatography (HPLC). The HPLC separations revealed that corticosterone was extensively metabolized mainly to more polar substances. Regarding the types of metabolites formed, significant differences were found between males and females, but not between the experiments. Additionally, the immunoreactivity of these metabolites was assessed by screening the HPLC fractions with four enzyme immunoassays (EIA). However, only a newly established EIA for 5alpha-pregnane-3beta,11beta,21-triol-20-one (measuring corticosterone metabolites with a 5alpha-3beta,11beta-diol structure) detected several peaks of radioactive metabolites with high intensity in both sexes, while the other EIAs showed only minor immunoreactivity. Thus, our study for the first time provides substantial information about metabolism and excretion of corticosterone in urine and feces of mice and is the first demonstrating a significant impact of the animals' sex and the time of day. Based on these data it should be possible to monitor adrenocortical activity non-invasively in this species by measuring fecal corticosterone metabolites with the newly developed EIA. Since mice are extensively used in research world-wide, this could open new perspectives in various fields from ecology to behavioral endocrinology.
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