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Abbruzzetti, S., Viappiani, C., Small, J. R., Libertini, L. J., & Small, E. W. (2001). Kinetics of histidine deligation from the heme in GuHCl-unfolded Fe(III) cytochrome C studied by a laser-induced pH-jump technique. J Am Chem Soc, 123(27), 6649–6653.
Abstract: We have developed an instrumental setup that uses transient absorption to monitor protein folding/unfolding processes following a laser-induced, ultrafast release of protons from o-nitrobenzaldehyde. The resulting increase in [H(+)], which can be more than 100 microM, is complete within a few nanoseconds. The increase in [H(+)] lowers the pH of the solution from neutrality to approximately 4 at the highest laser pulse energy used. Protein structural rearrangements can be followed by transient absorption, with kinetic monitoring over a broad time range (approximately 10 ns to 500 ms). Using this pH-jump/transient absorption technique, we have examined the dissociation kinetics of non-native axial heme ligands (either histidine His26 or His33) in GuHCl-unfolded Fe(III) cytochrome c (cyt c). Deligation of the non-native ligands following the acidic pH-jump occurs as a biexponential process with different pre-exponential factors. The pre-exponential factors markedly depend on the extent of the pH-jump, as expected from differences in the pK(a) values of His26 and His33. The two lifetimes were found to depend on temperature but were not functions of either the magnitude of the pH-jump or the pre-pulse pH of the solution. The activation energies of the deligation processes support the suggestion that GuHCl-unfolded cyt c structures with non-native histidine axial ligands represent kinetic traps in unfolding.
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Wolter, R., Pantel, N., Stefanski, V., Möstl, E., & Krueger, K. (2014). The role of an alpha animal in changing environmental conditions. Physiol. Behav., 133, 236–243.
Abstract: Abstract The maintenance and development of conservation areas by grazing of large herbivores, such as Przewalski's horses, is common practice. Several nature conservation areas house male bachelor groups of this species. When males are needed for breeding they are removed from the groups, often without considering group compositions and individual social positions. However, alpha animals are needed for ensuring group stability and decision making in potentially dangerous situations in several species. To investigate the role of the alpha male in a bachelor group, we observed the behaviour of five Przewalski's horse males during the enlargement of their enclosure. We analyzed the group's social structure and movement orders, as well as the animals' connectedness, activity budgets, and whether they moved with preferred group members and how factors such as social rank influenced the horses' behaviour. We also investigated the excretion of glucocorticoid metabolites (GCM) via faeces of the horses while exploring a new area as a parameter of glucocorticoid production. Our results show that the alpha male is important for a bachelor group in changing environmental conditions. The alpha male had the highest level of connectedness within the group. When exploring the new environment, its position in the group changed from previously being the last to being the first. Furthermore the whole group behaviour changed when exploring the new area. The stallions showed reduced resting behavior, increased feeding and did not stay close to each other. We found that the excretion of glucocorticoid metabolites of most horses rose only marginally during the first days on the new area while only the alpha male showed a significant increased amount of glucocorticoid production during the first day of the enclosure enlargement.
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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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Hartmann, E., Søndergaard, E., & Keeling, L. J. (2012). Identifying potential risk situations for humans when removing horses from groups. Appl. Anim. Behav. Sci., 136(1), 37–43.
Abstract: Removing a horse from its social group may be considered risky, both for the handler and the horse, because other horses can interfere in the catching process. The main aim of this study was to identify where and when these risk situations occur while removing a horse from its group. A potential risk situation was defined by the closeness of loose horses in the group or by any physical contact with them. Whether the number of horses following would be influenced by the social rank of the horse being led out, and whether more horses would follow to the gate when a larger proportion of the group was removed compared to when a single horse was taken out were also investigated. Thirty-two mares (1–2 years) were kept in groups of four. All horses were taken out of their home paddock twice alone (64 tests) and twice with a companion (32 tests). One handler (or two handlers when two horses were removed) was asked to approach (phase 1) and catch the target horse (phase 2), walk it to the centre of the paddock and remain stationary at a post for 30 s (phase 3), walk to the paddock entrance (phase 4) and through the gate (phase 5). The number of horses following, and the number of loose horses in proximity (<2 m, 2–5 m) to the target horse and handler was estimated, and horse–horse and horse–human interactions were recorded continuously for the five scoring phases. Significantly more loose horses were within 2 m of a single target horse during the phases approach (mean ± SD: 1.5 ± 0.8), catch (1.6 ± 0.9) and post (1.7 ± 0.7) than during walk (1.0 ± 0.5) and gate (1.1 ± 0.6). Rank did not influence the number of horses following to the gate (high rank: 2.4 ± 0.7; lower rank: 2.0 ± 1.0; P = 0.396) and interactions between horses were rare. A greater proportion of the loose horses followed when two horses (0.9 ± 0.2) were removed compared to when a single horse (0.7 ± 0.3) was taken out (P = 0.011). In conclusion, maintaining a distance to other horses in the group by reducing the time being relatively stationary, so giving loose horses fewer chances to approach, is likely to contribute to improved handler's safety. Removing a small proportion of the group may also decrease the probability of the other horses following.
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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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Meunier, H., Leca, J. B., Deneubourg, J. L., & Petit, O. (2006). Group movement decisions in capuchin monkeys: the utility of an experimental study and a mathematical model to explore the relationship between individual and collective behaviours. Behaviour, 143, 1511–1527.
Abstract: In primate groups, collective movements are typically described as processes dependent on leadership mechanisms. However, in some species, decision-making includes negotiations and distributed leadership. These facts suggest that simple underlying processes may explain certain decision mechanisms during collective movements. To study such processes, we have designed experiments on white-faced capuchin monkeys (Cebus capucinus) during which we provoked collective movements involving a binary choice. These experiments enabled us to analyse the spatial decisions of individuals in the group. We found that the underlying process includes anonymous mimetism, which means that each individual may influence all members of the group. To support this result, we created a mathematical model issued from our experimental data. A totally anonymous model does not fit perfectly with our experimental distribution. A more individualised model, which takes into account the specific behaviour of social peripheral individuals, revealed the validity of the mimetism hypothesis. Even though white-faced capuchins have complex cognitive abilities, a coexistence of anonymous and social mechanisms appears to influence their choice of direction during collective movements. The present approach may offer vital insights into the relationships between individual behaviours and their emergent collective acts.
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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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Hinde, R. A. (1969). Analyzing the roles of the partners in a behavioral interaction--mother-infant relations in rhesus macaques. Ann N Y Acad Sci, 159(3), 651–667.
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Conradt, L., Krause, J., Couzin, I. D., & Roper, T. J. (2009). “Leading According to Need” in Self-Organizing Groups. Am Nat, 173(3), 304–312.
Abstract: Self‐organizing‐system approaches have shed significant light on the mechanisms underlying synchronized movements by large groups of animals, such as shoals of fish, flocks of birds, or herds of ungulates. However, these approaches rarely consider conflicts of interest between group members, although there is reason to suppose that such conflicts are commonplace. Here, we demonstrate that, where conflicts exist, individual members of self‐organizing groups can, in principle, increase their influence on group movement destination by strategically changing simple behavioral parameters (namely, movement speed, assertiveness, and social attraction range). However, they do so at the expense of an increased risk of group fragmentation and a decrease in movement efficiency. We argue that the resulting trade‐offs faced by each group member render it likely that group movements are led by those members for which reaching a particular destination is most crucial or group cohesion is least important. We term this phenomenon leading according to “need” or “social indifference,” respectively. Both kinds of leading can occur in the absence of knowledge of or communication about the needs of other group members and without the assumption of altruistic cooperation. We discuss our findings in the light of observations on fish and other vertebrates.
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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.
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