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Suagee-Bedore, J. K., Linden, D. R., & Bennett-Wimbush, K. (2021). Effect of Pen Size on Stress Responses of Stall-Housed Horses Receiving One Hour of Daily Turnout. J. Equine Vet. Sci., 98, 103366.
Abstract: 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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Burla, J. B., Rufener, C., Bachmann, I., Gygax, L., Patt, A., & Hillmann, E. (2015). Effect of varying dimensions of the littered lying area on the lying behaviour of group-housed horses (Equus ferus caballus). In Proceedings of the 3. International Equine Science Meeting.
Abstract: Although horses can sleep while standing, recumbency is required for REM sleep and since all sleep stages must be completed for an entire sleep cycle, the opportunity for recumbency is essential for animal welfare. Observations on feral horses indicate a minimal lying duration of 30 min, preferably on a deformable and dry ground. In contrast to feral horses, lying behaviour in stabled horses is often affected by the dimensions of the provided lying area and rank.
In Switzerland, minimum requirements (MR) for the littered lying area are established by law to ensure animal welfare (BLV, 2008) (A/N: approximately match German recommendations (BMEL, 2009)). The aim of this study was to assess the adequacy of the dimensions of the minimum requirements for group-housed horses by investigating 38 horses in 8 groups. Further, hard rubber mats were provided supplementary in order to assess their suitability as an alternative to litter. Four treatments were each applied in randomised order:
– 0x MR: no litter + 1.5x MR with rubber mats
– 0.5x MR: 0.5x MR with litter + 1x MR with rubber mats
– 1x MR: 1x MR with litter; 0.5x MR with rubber mats
– 1.5x MR: 1.5x MR with litter + no rubber mats
For each treatment, after a habituation period of 8 days, lying behaviour was recorded (video, accelerometers) continuously for 72 hrs. Statistical analysis was performed using mixed effects models.
Regardless of the ground chosen, the duration of recumbency per 24 hrs was increasing with increasing dimensions of the littered area (F1,93 = 12.9, p = 0.0005; Fig. 1). Whereas the effect flattened from 1x to 1.5x MR, the duration spent on litter – a deformable ground – was increasing continuously (F1,62 = 23.1, p < 0.0001). Further, the proportion of lateral recumbency was increased with increased dimensions of the littered area (F1,79 = 12.3, p = 0.0007). Regarding the number of lying bouts, no differences were apparent between treatments providing litter, but recumbency occurred very seldom if only rubber mats were provided (F1,93 = 14.7, p = 0.0002). Further, low-ranking horses spent more lying bouts on rubber mats than high-ranking horses (F1,29 = 4.4, p = 0.04). Additionally, the larger the dimensions of the littered area the more horses were present in the lying area at the moment of lying down (F1,79 = 6.6, p = 0.01). Moreover, low-ranking horses showed considerably higher percentages of involuntarily terminated lying bouts than high-ranking horses if 0.5x and 1x MR were littered (F1,76 = 8.43, p = 0.005).
Although the indicated minimal lying duration was averagely performed, large individual differences occurred and at least 8% were lying down less than 30 min per 24 hrs in every treatment. Further, the inclusion of social parameters indicated a beneficial effect of an exceedance of the minimum requirements especially for low-ranking horses. Therefore, the minimum requirements established by Swiss law can be stated as adequate but should be perceived as minimum and not optimum dimensions.
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Smith, L. A., Wells, K. L., Marion, G., Swain, D. L., & Hutchings, M. R. (). Effects of group composition on the grazing behaviour of herbivores. Anim. Behav., In Press, Corrected Proof.
Abstract: Animal behaviour is often a function of the animal's physiological state. Groups of animals will often contain individuals with a range of physiological states and the grazing behaviour of herbivores is affected by their physiological state. This study compared the grazing decisions of animals in groups of single and mixed physiological states. Using a grazing model that simulated individual herbivore behaviour in relation to environmental distributions of forage resource (grass) and parasites (faeces), we tested the hypothesis that an animal's level of parasite exposure via the faecal-oral route is affected by the composition of physiological states in the group. Four physiological states were considered: parasite-naïve, parasitized, lactating and parasite-immune animals. Baseline parasite exposure levels for each state were generated by simulating single-state groups and were compared to simulations of each of the six two-state combinations. In single-state groups parasitized animals had the least and lactating animals had the greatest levels of parasite exposure. When co-grazing with lactating animals, parasitized, immune and naïve animals increased their parasite exposure, relative to single-state groups. When co-grazing with parasitized animals, lactating, immune and naïve animals reduced their parasite exposure, relative to single-state groups. There was no difference in parasite exposure of the immune or naïve animals co-grazing together when compared to the single-state groups. These results highlight the need to recognize the impact of the individual when studying group-living animals.
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Christensen, J. W., Ladewig, J., Sondergaard, E., & Malmkvist, J. (2002). Effects of individual versus group stabling on social behaviour in domestic stallions. Appl. Anim. Behav. Sci., 75(3), 233–248.
Abstract: Domestic horses (Equus caballus) are typically kept in individual housing systems, in which they are deprived of physical contact. In order to study the effects of social restrictions on behaviour in young horses, nineteen 2-year-old stallions were housed either singly (n=7), or in groups of three (n=12) for 9 months. Subsequently, the stallions were released into two separate 2 ha enclosures according to treatment, and recordings were made on social interactions and nearest neighbours during a 6-week-period, 28 h per week. Previously group stabled stallions frequently had a former group mate as their nearest neighbour (P=0.001), whereas previously singly stabled stallions did not associate more with their former box neighbours, to whom physical contact was limited by bars during the previous treatment. The nearest neighbour was more frequently recorded to be within one horselength of singly stabled than of group stabled stallions (P=0.005). More aggressive behaviour was recorded in the group of previously singly stabled stallions, i.e. bite threats (P=0.032), whereas group stabled stallions tended to make more use of subtle agonistic interactions (displacements, submissive behaviour). Singly stabled stallions also responded to the 9 months of social deprivation by significantly increasing the level of social grooming (P<0.001) and play behaviour (P<0.001), when subsequently interacting freely with other horses. The increased occurrence may relate to a build-up of motivation (a rebound effect), as well as to external factors, such as playful pasture companions and the increased space allowance of the pasture. It is concluded that 2-year-old domestic stallions are sensitive to social deprivation and that stabling has long-term effects, lasting 6 weeks at least, on the social behaviour in stallions.
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Suzuki, Y., & Toquenaga, Y. (2005). Effects of information and group structure on evolution of altruism: analysis of two-score model by covariance and contextual analyses. J. Theor. Biol., 232(2), 191–201.
Abstract: An altruistic individual has to gamble on cooperation to a stranger because it does not know whether the stranger is trustworthy before direct interaction. Nowak and Sigmund (Nature 393 (1998a) 573; J. Theor. Biol. 194 (1998b) 561) presented a new theoretical framework of indirect reciprocal altruism by image scoring game where all individuals are informed about a partner's behavior from its image score without direct interaction. Interestingly, in a simplified version of the image scoring game, the evolutionarily stability condition for altruism became a similar form of Hamilton's rule, i.e. inequality that the probability of getting correct information is more than the ratio of cost to benefit. Since the Hamilton's rule was derived by evolutionarily stable analysis, the evolutionary meaning of the probability of getting correct information has not been clearly examined in terms of kin and group selection. In this study, we applied covariance analysis to the two-score model for deriving the Hamilton's rule. We confirmed that the probability of getting correct information was proportional to the bias of altruistic interactions caused by using information about a partner's image score. The Hamilton's rule was dependent on the number of game bouts even though the information reduced the risk of cooperation to selfish one at the first encounter. In addition, we incorporated group structure to the two-score model to examine whether the probability of getting correct information affect selection for altruism by group selection. We calculated a Hamilton's rule of group selection by contextual analysis. Group selection is very effective when either the probability of getting correct information or that of future interaction, or both are low. The two Hamilton's rules derived by covariance and contextual analyses demonstrated the effects of information and group structure on the evolution of altruism. We inferred that information about a partner's behavior and group structure can produce flexible pathways for the evolution of altruism.
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Christensen, J. W., Søndergaard, E., Thodberg, K., & Halekoh, U. (2011). Effects of repeated regrouping on horse behaviour and injuries. Appl. Anim. Behav. Sci., 133(3), 199–206.
Abstract: Domestic horses are faced with social challenges throughout their lives due to limitations in social contact, space restrictions and frequent changes in social companionship. This is in contrast to natural conditions where horses live in relatively stable harem bands. Currently, little is known about how repeated regrouping affect horse behaviour and welfare, and it is unknown whether horses may adapt to regrouping. In this study, we aimed to investigate the effects of an unstable group structure, caused by weekly regroupings, on behaviour and frequency of injuries in young horses. Forty-five horses were included in the study and were randomly assigned to the treatments; Stable (S; seven groups of three horses) or Unstable (U; eight groups of three horses). The experimental period lasted 7 weeks, during which horses in Stable groups remained in the same group, whereas one horse was exchanged between Unstable groups every week. The groups were kept in 80m×80m grass-covered enclosures and were fed additional roughage on the ground daily. Social interactions were recorded in Unstable groups immediately after each regrouping (30min), and in both Stable and Unstable groups on day 1, 3 and 6 after each regrouping (2×20min/group/day). Injuries were scored by the end of the experimental period. The level of aggression shown by horses in Unstable groups immediately after regrouping was not affected by week (F5,35=0.42, P=0.83), indicating that horses neither habituated, nor sensitized, to repeated regrouping. Compared to horses in Stable groups, more agonistic behaviour was shown by horses in Unstable groups (i.e. non-contact agonistic; F1,65=5.60, P=0.02), whereas there was no treatment effect on other variables. The level of play behaviour appeared, however, to be more variable in Unstable groups. There was a significant effect of week on the level of contact agonistic interactions as well as greeting behaviour, due to a high occurrence in weeks 4-6. Non-contact agonistic interactions constituted the major part of agonistic interactions (66%). Possibly as consequence, no serious injuries were registered and there was no treatment effect (U=184; P=0.11). We conclude that the behaviour of young horses is affected by group management, and that horses appear not to adapt to weekly regroupings.
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Wilson, M. T., Silvestrini, M. C., Morpurgo, L., & Brunori, M. (1979). Electron transfer kinetics between Rhus vernicifera stellacyanin and cytochrome c (horse heart cytochrome c and Pseudomonas cytochrome c551). J Inorg Biochem, 11(2), 95–100.
Abstract: The electron transfer reactions between Rhus vernicifera stellacyanin and either horse heart cytochrome c or Pseudomonas aeruginosa cytochrome c551 were investigated by rapid reaction techniques. The time course of electron transfer is monophasic under all conditions, and thus consistent with a simple formulation of the reaction. Both stopped-flow and temperature-jump experiments yield equilibrium constants in reasonable agreement with values calculated from the redox potentials. The differences in reaction rate between the two cytochromes and stellacyanin are discussed in terms of the Marcus theory.
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Fureix, C., Bourjade, M., Henry, S., Sankey, C., & Hausberger, M. (2012). Exploring aggression regulation in managed groups of horses Equus caballus. Appl. Anim. Behav. Sci., 138(3–4), 216–228.
Abstract: Horses are highly social animals that have evolved to live in social groups. However, in modern husbandry systems, single housing prevails where horses experience social isolation, a challenge-to-welfare factor. One major reason for this single housing is the owners’ concerns that horses may injure each other during aggressive encounters. However, in natural conditions, serious injuries due to aggressive encounters are rare. What could therefore explain the claimed risks of group living for domestic horses? Basing our questioning on the current knowledge of the social life of horses in natural conditions, we review different practices that may lead to higher levels of aggression in horses and propose practical solutions. Observations of natural and feral horses mostly indicate a predominance of low frequencies and mild forms of aggression, based on subtle communication signals and ritualized displays and made possible by group stability (i.e. stable composition), dominance hierarchy and learning of appropriate social skills by young horses. Obviously, adults play a major role here in canalizing undesirable behaviours, and social experience during development, associated with a diversity of social partners, seems to be a prerequisite for the young horse to become socially skilled. Given the natural propensity of horses to have a regulation of aggression in groups, the tendency to display more aggression in groups of domestic horses under some management practices seems clearly related to the conditions offered. We therefore review the managing practices that could trigger aggressiveness in horses. Non social practices (space, resource availability) and social practices (group size, stability of membership, composition and opportunities for social experiences during development) in groups of domestic horses are discussed here. Finally, we propose simple practical solutions leading to more peaceful interactions in groups of domestic horses, based on the knowledge of horses’ natural social life which therefore should be enhanced (e.g. ensuring roughage availability, favouring group stability, introducing socially experienced adults in groups of young horses, etc.). The state of the art indicates that many questions still need to be answered. Given the importance of the associated welfare issues and the consequences on the use of horses, further research is required, which could benefit horses… and humans.
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Knopff, K., & Pavelka, M. (2006). Feeding Competition and Group Size in Alouatta pigra. Int. J. Primatol., 27(4), 1059–1078.
Abstract: Researchers consider group size in primates to be determined by complex relationships among numerous ecological forces. Antipredator benefits and better resource defense are the primary pressures for large groups. Conversely, intragroup limited food availability, can result in greater intragroup feeding competition and individual energy expenditure in larger groups, creating energetic advantages for individuals in small groups and placing an upper limit group size. However, the extent to which food availability constrains group size remains unclear for many species, including black howlers (Alouatta pigra), which ubiquitously live in small social groups (≤10 individuals). We studied the relationship between group size and 2 key indices of feeding competition-day journey length and activity budgets in 3 groups of wild Alouatta pigra at a hurricane-damaged site in Belize, Central America. We controlled for differences in food availability between home ranges (food tree density) and compared both indicators of feeding competition directly with temporal variation in food availability for each group. Our results show no consistent association between resource availability, group size, and either index of competition, indicating that feeding competition does not limit group size at the site i.e., that larger groups can form without increased costs of feeding competition. The results support the search for other explanations, possibly social ones, for small group size in the primates, and we conclude with suggestions and evidence for such alternative explanations.
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Hoang, L., Maity, H., Krishna, M. M. G., Lin, Y., & Englander, S. W. (2003). Folding units govern the cytochrome c alkaline transition. J Mol Biol, 331(1), 37–43.
Abstract: The alkaline transition of cytochrome c is a model for protein structural switching in which the normal heme ligand is replaced by another group. Stopped flow data following a jump to high pH detect two slow kinetic phases, suggesting two rate-limiting structure changes. Results described here indicate that these events are controlled by the same structural unfolding reactions that account for the first two steps in the reversible unfolding pathway of cytochrome c. These and other results show that the cooperative folding-unfolding behavior of protein foldons can account for a variety of functional activities in addition to determining folding pathways.
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