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Wilson, M.T.; Silvestrini, M.C.; Morpurgo, L.; Brunori, M. |
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Electron transfer kinetics between Rhus vernicifera stellacyanin and cytochrome c (horse heart cytochrome c and Pseudomonas cytochrome c551) |
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1979 |
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Journal of Inorganic Biochemistry |
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J Inorg Biochem |
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11 |
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2 |
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95-100 |
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Animals; Copper; Cytochrome c Group/*metabolism; Electron Transport; Kinetics; Metalloproteins/*metabolism; Plant Proteins/*metabolism; *Plants, Toxic; Pseudomonas aeruginosa/*metabolism; Toxicodendron/*metabolism |
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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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0162-0134 |
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PMID:228006 |
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3879 |
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Tsong, T.Y. |
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Conformational relaxations of urea- and guanidine hydrochloride-unfolded ferricytochrome c |
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Journal Article |
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1977 |
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The Journal of Biological Chemistry |
Abbreviated Journal |
J Biol Chem |
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252 |
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24 |
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8778-8780 |
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*Cytochrome c Group; Guanidines/*pharmacology; Protein Conformation/drug effects; Spectrometry, Fluorescence; Urea/*pharmacology |
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Several recent studies of protein the unfolded proteins. In urea- and guanidine HCl-unfolded ferricytochrome c (horse heart), an acid-induced spin state transformation of the heme group has been detected by the heme absorptions, Trp-59 fluorescence, and the intrinsic viscosity of protein. Kinetics of this second conformational transition, by the temperature jump and stopped flow methods, are complex. One rapid reaction (tau1), pH-independent, occurs in a 50-mus range; the second reaction (tau2), in a 1-ms range, depends linearly upon pH and is faster at the alkaline side; a third reaction (tau3), in a 1-s range, shows a sigmoidal transition at pH 5.1 and is faster at the acidic side. The results are consistent with a kinetic scheme which involves protein conformational changes in the transformation of the heme coordination state. The kinetics, along with previous equilibrium studies, indicate that ligand or charge interactions within a protein molecule are not completely prohibited even in strongly denaturing conditions, such as in high concentrations of urea and guanidine HCl. Thus, local structures of peptide chain associated with these interactions can exist in the unfolded protein. |
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0021-9258 |
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PMID:200618 |
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refbase @ user @ |
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3882 |
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Fischhoff, I.R.; Sundaresan, S.R.; Cordingley, J.; Larkin, H.M.; Sellier, M.-J.; Rubenstein, D.I. |
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Social relationships and reproductive state influence leadership roles in movements of plains zebra, Equus burchellii |
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Journal Article |
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Year |
2007 |
Publication |
Animal Behaviour. |
Abbreviated Journal |
Anim. Behav. |
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73 |
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5 |
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825-831 |
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collective movements; drinking; equid; Equus burchellii; foraging; group dynamics; motivation; plains zebra; reproductive status; social relationships |
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In animal groups, collective movements emerge from individual interactions. Biologists seek to identify how characteristics of actors in these groups, and their relationships, influence the decision-making process. We distinguished two basic factors determining leadership in group choices: identity and state. We hypothesized that identity is more important to leadership in groups with stable relationships, which permit the development of habitual roles. In groups with fluid membership, particular individuals or subgroups are less likely to emerge as consistent leaders. Instead, we predicted that movement initiation in unstable groups depends on individual state at the time of the decision. We characterized how identity and reproductive state influenced leadership patterns in the movements of plains zebra. As in many other mammals, lactation in this species significantly alters water and energy needs. We investigated leadership in tightly knit harems and loosely bonded herds of multiple harems. Harem females tended to have habitual roles in the initiation of harem movement. In herds, however, we found no consistent leaders among harems. At both levels of social organization, lactation was a key determinant of leadership. In harems, lactating females were more likely to initiate movement than nonlactating females. In turn, harems containing lactating females were more likely to lead herd movements. Thus, we conclude that social relationships and reproductive state together shape the interactions that produce group behaviours. One benefit to lactating females of leading herd movements is preferential access to scarce water. Thus, leadership roles in group decisions may have fitness consequences. |
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Equine Behaviour @ team @ room B 3.092 |
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1825 |
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Levin, L.E. |
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Passage order through different pathways in groups of schooling fish, and the diversified leadership hypothesis |
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1996 |
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Behavioural Processes |
Abbreviated Journal |
Behav. Process. |
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37 |
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1 |
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1-8 |
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Animal sociality; Inter-individual variability; Aggregation-dispersion; Group problem solving |
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The diversified leadership hypothesis proposes that different individuals within a school of fish act as leaders in different circumstances. This `circumstantial leadership' results from inter-individual behavioral variability and a `cohesion-dispersion' tendency modulated by `failure-success' contingencies. The hypothesis predicts that when offered different pathways to escape the restriction of their swimming space, individuals within a group of fish will show 1. (a) consistent passage orders in each pathway, but2. (b) different passage orders in different pathways. Using an avoidance paddle and three different groups of fish (Aphyocharax erithrurus) the results confirmed prediction 1. (a) while prediction2. (b) was verified only in one group. |
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Equine Behaviour @ team @ room B 3.029 |
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2069 |
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Author |
Flauger, B. |
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Title |
The introduction of horses into new social groups with special regard to their stress level |
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2011 |
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Pferd; Equiden; Eingliederungstechnik; Integrationspferd; Stress; Cortisol; Endokrine Reaktion; Gruppenhaltung; Verletzungsgefahr; Aggression; Futterplatzwahl; Kot; Geruchssinn; Mensch-Pferd Interaktion; horse; equids; introduction technique; integration horse; stress; cortisol; endocrine response; group housing; injury risk; aggression; feeding decision; faecal sample; olfaction; human-horse interaction |
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Horses are a highly social species living in complex social systems which should require them to memorise and generalise social experiences and distinguish between familiar and unfamiliar conspecifics. In the main part of my thesis I concentrated on the specific conflict situation of a horse being introduced into a new social group, and investigated its behaviour and stress level. Horses were either introduced (1) immediately, (2) after an observation period, or (3) together with an integration horse after an observation period. Additionally, in the second part of my thesis I arranged several experiments to elaborate additional aspects which could affect the behaviour of horses during introductions. In this study I could describe a simplified method for measuring stress through the analysis of faecal GCMs in horses. An enzyme immunoassay (EIA) for 11-oxoaetiocholanolone using 11-oxoaetiocholanolone-17-CMO: BSA (3?,11-oxo-A EIA) as antigen showed high amounts of immunoreactive substances. The new assay increases the accuracy of the test and lowers the expenses per sample; also storing of samples at room temperature after collection is less critical. This is a big advantage both in the field of wildlife management of equids and in the field of equestrian sports (chapter 1). Comparing the different introduction techniques, the introduction with an integration horse led to significantly less total interactions and lower levels of aggression than the introduction of single horses, both immediately and after several days of observing the new group. Additionally, by observing the behaviour of the horses during everyday sociality I could develop a formula describing the interrelationship between expected aggression level and enclosure size per horse. The curve takes an exponential shape. Starting from a space allowance of 300 m2 and more per horse, the amount of aggressions per hour approaches zero. For the reduction of aggression levels and injury risks in socially kept horses I recommend an enclosure size of at least 300 m2 per horse (chapter 2). I further investigated the stress level of the introduced animals. Horses which were immediately introduced did not show elevated faecal GCMs. In contrast, horses which were introduced after an observation period had slightly elevated values 2 and 3 days after the introduction. For horses introduced together with an integration horse faecal GCMs were significantly above the baseline value on the day of introduction and 1 day after it. These differences between introduction techniques indicate that the introduction event itself is not as stressful as previously assumed. Rather standing together with an integration horse and not being able to integrate immediately into the complete group elicits stress in horses (chapter 3). In the commentary of chapter 4 several studies are discussed which failed to demonstrate social learning in horses. It is argued that they did not consider important aspects which could have an influence, such as the dominance status or the social background of the horses (chapter 4). In chapter 5 a social feeding situation was investigated. The social rank as well as the position of conspecifics affected the feeding strategy of horses. Domestic horses used social cognition and strategic decision making in order to decide where to feed. When possible they tended to return to the same, continuously supplied feeding site and switched to an ?avoidance tendency? in the presence of dominant horses or when another horse was already feeding there (chapter 5). One possibility to recognize group members is through olfactory recognition. In chapter 6 it is shown that horses are able to distinguish their own from their conspecifics? faeces. In addition, they paid most attention to the faeces of those group members from which they received the highest amount of aggressive behaviour (chapter 6). Horses show cognitive abilities because they are able to use humans as local enhancement cues when searching for food, independently of their body posture or gaze consistency when the persons face them. Moreover, they seem to orientate on the attention of familiar persons more than of unfamiliar persons (chapter 7). Altogether, the results of this thesis provide further support for the view that horses show good conflict resolution strategies. They are perfectly able to deal with the conflict situation of being introduced to new group members, and the introduction event itself is not as stressful as previously assumed. It is rather suggested that standing together with an integration horse and not being able to integrate immediately into the complete group elicits stress in horses. All additional experimental set-ups could demonstrate that horses are well capable of social cognition. |
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Ph.D. thesis |
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Equine Behaviour @ team @ epub18463 |
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5736 |
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Author |
Dugatkin, L.A.; Godin, G.J. |
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Predator inspection, shoaling and foraging under predation hazard in the Trinidadian guppy,Poecilia reticulata |
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Journal Article |
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1992 |
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Environmental Biology of Fishes |
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34 |
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3 |
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265-276 |
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Antipredation – Social group – Feeding – Predation risk – Trade-off – Fish |
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Guppies,Poecilia reticulata, living in stream pools in Trinidad, West Indies, approached a potential fish predator (a cichlid fish model) in a tentative, saltatory manner, mainly as singletons or in pairs. Such behavior is referred to as predator inspection behavior. Inspectors approached the trunk and tail of the predator model more frequently, more closely and in larger groups than they approached the predator's head, which is presumably the most dangerous area around the predator. However, guppies were not observed in significantly larger shoals in the stream when the predator model was present. In a stream enclosure, guppies inspected the predator model more frequently when it was stationary compared to when it was moving, and made closer inspections to the posterior regions of the predator than to its head. Therefore, the guppies apparently regarded the predator model as a potential threat and modified their behavior accordingly when inspecting it. Guppies exhibited a lower feeding rate in the presence of the predator, suggesting a trade-off between foraging gains and safety against predation. Our results further suggest that predator inspection behavior may account for some of this reduction in foraging. These findings are discussed in the context of the benefits and costs of predator inspection behavior. |
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Equine Behaviour @ team @ |
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2176 |
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Nuñez, C.M.V.; Adelman, J.S.; Smith, J.; Gesquiere, L.R.; Rubenstein, D.I. |
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Linking social environment and stress physiology in feral mares (Equus caballus): Group transfers elevate fecal cortisol levels |
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2014 |
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General and Comparative Endocrinology |
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196 |
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26-33 |
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Equus caballus; Fecal cortisol; Feral mare; Group transfer; Stress; Social instability |
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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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0016-6480 |
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Equine Behaviour @ team @ |
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5743 |
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Christensen, J.W.; Ladewig, J.; Sondergaard, E.; Malmkvist, J. |
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Effects of individual versus group stabling on social behaviour in domestic stallions |
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2002 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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75 |
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3 |
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233-248 |
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Horse; Individual stabling; Group stabling; Social behaviour |
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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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Equine Behaviour @ team @ |
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2257 |
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Le Pendu, Y.; Guilhem, C.; Briedermann, L.; Maublanc, M.-L.; Gerard, J.-F. |
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Interactions and associations between age and sex classes in mouflon sheep (Ovis gmelini) during winter |
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2000 |
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Behavioural Processes |
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Behav. Process. |
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52 |
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2-3 |
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97-107 |
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Group composition; Interactive behaviour; Sexual segregation; Social organisation; Ungulate; Wild sheep |
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Equine Behaviour @ team @ |
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4248 |
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VanDierendonck, M.C. |
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Social relationships in a group of horses without a mature stallion |
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2006 |
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The Importance of Social Relationships in Horses |
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Chapter 4 |
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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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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. |
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Universiteit Utrecht |
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Equine Behaviour @ team @ |
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2369 |
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