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Goursot, C., Düpjan, S., Puppe, B., & Leliveld, L. M. C. (2021). Affective styles and emotional lateralization: A promising framework for animal welfare research. Appl. Anim. Behav. Sci., 237, 105279.
Abstract: The growing recognition of animals as individuals has broader implications for farm animal welfare research. Even under highly standardized on-farm conditions, farm animals show heterogeneous but individually consistent behavioural patterns towards various stimuli, based on how they appraise these stimuli. As a result, animal welfare is likely to be highly individual as well, and studying the proximate mechanisms underlying distinct individual behaviour patterns and appraisal will improve animal welfare research. We propose to extend the framework of affective styles to bridge the gap between existing research fields on animal personality and affective states. Affective styles refer to consistent individual differences in emotional reactivity and regulation and can be predicted by baseline cerebral lateralization. Likewise, animals with consistent left or right motor biases--a proxy measure of individual patterns in cerebral lateralization--have been shown to differ in their personality, emotional reactivity, motivational tendencies or coping styles. In this paper, we present the current knowledge of the links between laterality and stable individual traits in behaviour and affect in light of hypotheses on emotional lateralization. Within our suggested framework, we make recommendations on how to investigate affective styles in non-human animals and give practical examples. This approach has the potential to promote a science of affective styles in nonhuman animals and significantly advance research on animal welfare.
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Stomp, M., d'Ingeo, S., Henry, S., Cousillas, H., & Hausberger, M. (2021). Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model. Appl. Anim. Behav. Sci., 236, 105271.
Abstract: Assessing the animal welfare state is a challenge given the subjective individual cognitive and emotional processing involved. Electroencephalography (EEG) spectrum analysis has proved an ecologically valid recording situation to assess the link between brain processes and affective or cognitive states in humans: a higher slow wave/fast wave ratio has been associated with a positive internal state. In particular, a high production of theta power (3-8 Hz) has been related to positive emotions. On the other hand, it has been hypothesized that a left hemisphere (LH) dominance may be associated with a better welfare state. Here, we test the hypothesis that individual differences in the resting-state quantitative EEG power spectrum of adult horses (N = 18) and its lateralization pattern may reflect individual subjective perception of their conditions of life and welfare state. The results show clear individual differences in the proportions of the different waves and their inter-hemispheric distribution. Three different EEG power spectrum profiles were highlighted, from a bilateral predominance of theta waves in horses in a more positive welfare state to a bilateral predominance of beta waves in horses with clear expressions of compromised welfare. Interestingly, particular correlations were found between wave power activity and welfare parameters. We found a negative correlation between the number of stereotypic behaviours per hour and the median proportion of theta waves in the left hemisphere. and between the overall state (total chronic stress score) of welfare and gamma production in the right hemisphere (RH). These findings go along the hypothesis of a particular involvement of the left hemisphere for positive processing and of the right hemisphere for negative processing. However, the pattern of laterality did not appear as the most important feature here as both extreme clusters in terms of welfare showed bilateral predominance of one wave type. It is possible that hemispheric specialization makes more sense during acute emotion-inducing conditions rather than in this resting-state context (i.e. in absence of any high emotion-inducing stimulation), although the opposition gamma versus theta waves between both hemispheres in the horses with an intermediate welfare state is noticeable and intriguing. It seems that bilateral but also LH theta activity is a promising neurophysiological marker of good welfare in horses, while a bilateral or RH high production of gamma waves should alert about potential welfare alterations. Quantitative resting-state EEG power spectrum appears as a highly promising tool for exploring the brain processes involved in the subjective perception of chronic welfare, as a useful complementary tool for welfare assessment.
Keywords: Laterality; Electroencephalography; Theta wave; Welfare; Horses
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Christensen, J. W., Munk, R., Hawson, L., Palme, R., Larsen, T., Egenvall, A., et al. (2021). Rider effects on horses' conflict behaviour, rein tension, physiological measures and rideability scores. Appl. Anim. Behav. Sci., 234, 105184.
Abstract: Many breeding organisations include a subjective scoring of rideability by a professional rider into their evaluation of sports horses, but the consistency and reliability of the scoring system is debateable. The aim of this study was to investigate (i) whether professional riders agree in their scoring of rideability, and (ii) whether rideability scores are affected by rein tension, horse conflict behaviour, heart rate, and salivary cortisol, and (iii) whether riders induce different levels of conflict behaviour and physiological responses in the horses. Ten professional, female riders each rode 10 dressage horses (level M German scale; n = 100 combinations) through a standardised dressage test (10 min warm-up followed by a 4-min test) and subsequently scored the horses for rideability on the official 1-10 scale (1 = poor to 10 = excellent) from the Danish Riding Federation. Rein tension, horse heart rate, saliva cortisol and conflict behaviour were measured for each rider-horse pair. The riders were inconsistent in their scoring of rideability to the individual horses, e.g. scores for one of the horses ranged from 1 to 8. There was a significant effect of rider (P = 0.003) and the frequency of conflict behaviour (undesired head movements: P < 0.001, breaking the gait: P = 0.013, and other evasive behaviour: P = 0.032) on rideability scores, i.e. the more conflict behaviour the lower the score. There was no significant effect of rein tension and the physiological measures on rideability scores. However, there was a significant effect of rider on rein tension, horses' heart rate and increases in saliva cortisol concentrations and a tendency for some types of conflict behaviour, suggesting that some riders induced more discomfort in the horses. Future studies could help shed light on which elements of riding style are particularly important for sports horse welfare. In conclusion, this study found a large variation in rideability scores assigned to ten sports horses by ten professional riders. Rideability scores were dependent on the level of horse conflict behaviour, but not rein tension and physiological measures. Further studies are needed to improve the objectivity, consistency and reliability of rideability assessment of sports horses.
Keywords: Equitation science; Heart rate; Horse riding; Sport horse; Temperament; Training cues
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Lesimple, C., Reverchon-Billot, L., Galloux, P., Stomp, M., Boichot, L., Coste, C., et al. (2020). Free movement: A key for welfare improvement in sport horses? Appl. Anim. Behav. Sci., 225, 104972.
Abstract: Horses, and in particular sport horses, remain housed predominantly in single stalls. One of the main reported reasons is the fear that they will become agitated and injure themselves and thereby impair their performance if released in paddocks. The hour spent daily at work is also assumed to be sufficient to satisfy the horses' needs for locomotion. Growing scientific evidence shows that single stall housing has negative consequences on horses' welfare and that time for free movement is necessary. Our aim was to assess the feasibility of allowing sport horses used to staying permanently in their stall (except for 1 h riding/day) daily free time in a paddock and to evaluate its potential impact on their welfare using two behavioural reliable indicators (stereotypic behaviours and ear position), as well as selected blood parameters (blood cell count, oxytocin and serotonin concentrations). Two experiments were conducted on the same site. The first experiment evaluated sport horses' habituation to the novel situation of daily sessions in a paddock, and recorded welfare indicators in their stall before and during the experiment, comparing horses that were taken outdoors (experimental) and those that stayed in their stall (control). The second experiment evaluated the impact of this daily free time in a paddock on horses' welfare and its durability, focusing on positive indicators. Behavioural observations in paddocks showed that even horses that had never experienced free movement outside their stall habituated rapidly to this situation. The presence of hay in the paddock, may have speeded up habituation. Their restricted living conditions were associated with abnormalities in blood cell count that were not overcome during the time of daily paddock sessions but behavioural indicators showed that their welfare improved. In the second study, the experimental horses' welfare improved during the paddock release period, in particular their stereotypic behaviours decreased and oxytocin levels increased. No effects on serotonin concentrations could be evidenced. These effects were directly associated with being in paddock, as the indicators returned rapidly to their previous levels indicating compromised welfare when the paddock release sessions stopped. In conclusion, it can be recommended to release sport horses for free movement in paddocks as welfare is improved and subjective assessment by caretakers indicated minimal risks.
Keywords: Horse; Housing; Free movement; Horse welfare; Stereotypic behaviour; Oxytocin
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Stueckle, S., & Zinner, D. (2008). To follow or not to follow: decision making and leadership during the morning departure in chacma baboons. Anim. Behav., 75(6), 1995–2004.
Abstract: To benefit from group living, group members need to keep the group cohesive by coordinating time and direction of travelling. Self-organization and leadership are two means of coordination and two types of decision can be made on the group level: combined and consensus. We studied the initiation process of group movements during the morning departure of a group of chacma baboons, Papio hamadryas ursinus, from its sleeping site in De Hoop Nature Reserve, South Africa. Findings from other female-bonded primate groups led us to hypothesize that females should play a major role in the decision-making process. Approximately 75% of the adults made a start attempt, with 62 of 92 attempts being by males. There was no sex difference in the probability of being successful when initiating an attempt. Lactating females initiated fewer than pregnant or cycling females. Thus, at least for this group of chacma baboons, leadership appeared to be distributed and the decision about the timing of departure and travel direction seemed to be a partially shared consensus decision with adult males contributing more to the decision outcome, with a slightly more prominent role of the dominant male. Our results do not support the [`]leading females' hypothesis. No behavioural patterns that might serve as specialized signals leading to a more successful recruitment of other group members were observed. The departure process appeared to be coordinated merely through individuals setting an example by moving off.
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Lusseau, D., Whitehead, H., & Gero, S. (2008). Incorporating uncertainty into the study of animal social networks. Anim. Behav., 75(5), 1809–1815.
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Udell, M. A. R., Dorey, N. R., & Wynne, C. D. L. (2008). Wolves outperform dogs in following human social cues. Anim. Behav., 76(6), 1767–1773.
Abstract: Domestic dogs, Canis familiaris, have been shown capable of finding hidden food by following pointing gestures made with different parts of the human body. However, previous studies have reported that hand-reared wolves, C. lupus, fail to locate hidden food in response to similar points in the absence of extensive training. The failure of wolves to perform this task has led to the proposal that the ability to understand others' intentions is a derived character in dogs, not present in the ancestral population (wolves). Here we show that wolves, given the right rearing environment and daily interaction with humans, can use momentary distal human pointing cues to find food without training, whereas dogs tested outdoors and dogs at an animal shelter do not follow the same human points. In line with past studies, pet dogs tested indoors were successful in following these points. We also show that the reported failure of wolves in some past studies may be due to differences in the testing environment. Our findings indicate that domestication is not a prerequisite for human-like social cognition in canids, and show the need for additional research on the role of rearing conditions and environmental factors in the development of higher-level cognitive abilities.
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Cheney, D. L., & Seyfarth, R. M. (1986). The recognition of social alliances among vervet monkeys. Anim. Behav., 34, 1722–1731.
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Valderrabano-Ibarra, C., Brumon, I., & Drummond, H. (2007). Development of a linear dominance hierarchy in nestling birds. Anim. Behav., 74(6), 1705–1714.
Abstract: Theoreticians propose that trained winning and losing are important processes in creating linear animal dominance hierarchies, and experiments have shown that both processes can occur in animals, but their actual roles in creating natural hierarchies are unknown. We described agonism in 18 broods of three blue-footed boobies, Sula nebouxii, a species for which trained winning and losing have been demonstrated, to infer how these processes generate and maintain a natural hierarchy. Ranks in the linear hierarchy that emerged in every brood were initially assigned by asymmetries in age, size and maturity, which led to differences between broodmates in levels of expressed and received aggression and, consequently, to differences in the training of their aggressiveness and submissiveness. Later, ranks appeared to be maintained by the chicks' acquired aggressive and submissive tendencies combined with ongoing effects of persisting differences in size and maturity. Our results suggest that trained winning and trained losing are important in the construction of booby hierarchies but that these two axes of learning are largely independent. Increase in submissiveness occurs over a period of about 10-20 days, and the level of submissiveness reached varies with the amount of aggression received. After training, submissiveness is apparently maintained by a lower level of aggression and increasing use of threats. Threats become increasingly effective as chicks age, but are never as effective as attacks.
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Jonart, L. M., Hill, G. E., & Badyaev, A. V. (2007). Fighting ability and motivation: determinants of dominance and contest strategies in females of a passerine bird. Anim. Behav., 74(6), 1675–1681.
Abstract: The communication of aggressive motivation or fighting ability has important fitness consequences for competing animals. Selection should favour rapid and honest communication between opponents to settle dominance relationships while avoiding prolonged and intense fighting. We investigated factors that influence fighting strategies and contest outcomes in female house finches, Carpodacus mexicanus, specifically focusing on the following questions. (1) What social contexts trigger an aggressive response? (2) Does body size and condition contribute to female fighting ability? (3) Do contextual factors, such as mate presence, nest status, nest proximity, and site experience contribute to fighting motivation? (4) Does contest intensity and duration increase as the differences in fighting ability between opponents decrease? (5) What is the relative contribution of fighting ability and aggressive motivation to the outcome of a contest? We found that aggression was triggered most frequently by female intrusions in the vicinity of nest and by extrapair female intrusions on an established pair. Female fighting and contest outcomes were strongly influenced by body condition and body size, and females were more motivated to initiate fights and won more contests when their mates were present. Females at the later breeding stages and those fighting closer to their nests were dominant and won more fights compared to females at earlier breeding stages or further from their nests. Females initiated a greater proportion of contests against opponents with similar local familiarity and breeding history. Escalated and prolonged contests, while rare, occurred exclusively between females of the most similar body size and condition. When differences in body condition between opponents are not easily perceived, contestants might escalate contests for more reliable assessments of relative fighting ability. Finally, body condition was not a strong determinant of contest outcome in the contexts with easily assessed differences in the resource value (e.g. mate presence), but without these motivational differences, body condition was the ultimate determinant of contest outcomes.
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