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Nicol, C. J. (1995). The social transmission of information and behaviour. Appl. Anim. Behav. Sci., 44(2-4), 79–98.
Abstract: Social influences on established behaviour and on the acquisition of new information and behaviour are reviewed. Distinctions between social facilitation and contagious behaviour are drawn and suggestions for further research on contagious behaviour are made. Socially derived visual, olfactory and auditory cues are considered as important influences on behaviour and subsequent learning. The evidence supporting two potential mechanisms of social learning, i.e. stimulus enhancement followed by individual learning, and imitation, is reviewed in detail. It is argued that the functions of social learning are similarly heterogeneous and include motor skill acquisition, gathering of environmental information, and social conformity. Factors affecting the spread of socially acquired skills, including the social relationship between demonstrator and observer, are highlighted. Lastly, the few studies of social learning that have been conducted with domestic species are reviewed and potential applied goals that could stimulate further research in this area are suggested.
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Nicol, C. J. (2002). Equine learning: progress and suggestions for future research. Appl. Anim. Behav. Sci., 78(2-4), 193–208.
Abstract: Horses are well able to form classical and instrumental associations and so the focus of much recent research has been on the stimulus control of instrumental learning. Horses appear to discriminate using spatial cues more easily than other stimulus features, as indicated both by the speed of initial task acquisition and by the extent to which acquired discriminations can be reversed. Phenomena associated with discrimination learning in laboratory animals, including generalisation and peak shift, have been demonstrated in horses. However, the ability of horses to classify stimuli into categories is more controversial. Although there is some evidence that horses may be able to form categories based on similarities in the physical appearance of different stimuli, there is currently no evidence that they are able to develop abstract concepts. Their performance on social learning tasks has also been poor. Few correlations are observed between the learning ability of individual horses on different tasks, suggesting that it may not be possible to classify individual horses as `good' or `poor' learners. Better learning performance by horses that are naturally calm is probably due to reduced interference in the learning process. Correct handling procedures can lower reactivity levels in horses, and may facilitate learning in some circumstances. Future research on equine learning needs to take into account the complex nature of equine social interaction. Studies on the effects of stress on learning, and on social and spatial cognition, are also particularly needed.
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Newberry, R. C., & Swanson, J. C. (2008). Implications of breaking mother-young social bonds. Appl. Anim. Behav. Sci., 110(1-2), 3–23.
Abstract: Whereas mammalian mothers and young may retain long-term social affiliations in nature, the management of animals in captivity typically dictates that offspring are abruptly and permanently separated from their mothers at a relatively early age, often prior to the time of natural weaning. For animal breeders, this strategy can enhance the yield of offspring from a breeding population. Morbidity and mortality can also precipitate severance of mother-young bonds. Although it is recognized that early weaning provides nutritional challenges for the young, relatively little attention has been paid to the psychological consequences and long-term impacts of breaking the mother-young bond in non-human mammals. Furthermore, whereas great strides are being made in our understanding of the neurobiological and genetic underpinnings of social bonding, the mechanisms underlying the process of detachment following establishment of a mother-young bond remain relatively unexplored, although parallels can be drawn with processes involved in withdrawal from addictive substances. In this review, we outline mechanisms involved in social bonding. We consider the diversity in extent and duration of mother-young attachment across mammalian lineages and implications for predicting the outcome of severing ties between mothers and young at different times post-partum. We identify characteristics signalling emotional distress resulting from separation of mothers and young and discuss strategies for mitigating separation-induced distress. These include postponement of separation, ensuring high-quality maternal care of young prior to separation, providing bonded individuals with opportunities to separate voluntarily for brief periods prior to permanent separation, use of anti-suck devices prior to separation, allowing a period of partial (fence line) contact prior to full separation, providing substitutes for stimuli previously exchanged between mother and young, providing social buffers, gradual introduction to new housing arrangements, and pharmacological intervention. Areas for future research are proposed, including the use of functional neuroimaging technologies and functional genomics approaches, in combination with behavioural assessments of reinstatement motivation, individual recognition memory and long-term consequences of early separation, to shed further light on the nature of mother-young bonding and detachment in animals.
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Neveu, P. J. (1993). Brain Lateralization and Immunomodulation. Int J Neurosci, 70(1-2), 135–143.
Abstract: The two sides of the brain may be differently involved in the modulation of immune responses as demonstrated by lesional and behavioral approaches in rodents. Lesions of right or left neocortex induced opposite effects on various immune parameters including mitogen-induced lymphoproliferation, interleukin-2 production, macrophage activation or natural killer cell activity. This animal model, useful to elucidate whereby the brain and the immune system can communicate, appears to be suitable for studying the immune perturbations observed during stroke in humans. Brain asymmetry in modulation of immune reactivity may also be demonstrated in intact animal using a behavioral paradigm. The direction of a lateralized motor behavior ie paw preference in a food reaching task, correlated with an asymmetrical brain organization, was shown to be associated with lymphocyte reactivity, natural killer cell activity and auto-antibody production. The association between paw preference and immune reactivity in mice varies according to the immune parameters tested and is a sex-dependent phenomenon in which genetic background may be involved. The experimental models for investigating asymmetrical brain modulation of the immune system should be useful for studying several physiological, pathological and genetic aspects of neuroimmunomodulation.
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Neumann Inga D, Veenema Alexa H, & Beiderbeck Daniela I. (2010). Aggression and anxiety: social context and neurobiological links. Frontiers in Behavioral Neuroscience, 4(1).
Abstract: BACKGROUND: Psychopathologies such as anxiety- and depression-related disorders are often characterized by impaired social behaviours including excessive aggression and violence. Excessive aggression and violence likely develop as a consequence of generally disturbed emotional regulation, such as abnormally high or low levels of anxiety. This suggests an overlap between brain circuitries and neurochemical systems regulating aggression and anxiety. In this review, we will discuss different forms of male aggression, rodent models of excessive aggression, and neurobiological mechanisms underlying male aggression in the context of anxiety. We will summarize our attempts to establish an animal model of high and abnormal aggression using rats selected for high (HAB) versus low (LAB) anxiety-related behaviour. Briefly, male LAB rats and, to a lesser extent, male HAB rats show high and abnormal forms of aggression compared with non-selected (NAB) rats, making them a suitable animal model for studying excessive aggression in the context of extremes in innate anxiety. In addition, we will discuss differences in the activity of the hypothalamic-pituitary-adrenal axis, brain arginine vasopressin, and the serotonin systems, among others, which contribute to the distinct behavioural phenotypes related to aggression and anxiety. Further investigation of the neurobiological systems in animals with distinct anxiety phenotypes might provide valuable information about the link between excessive aggression and disturbed emotional regulation, which is essential for understanding the social and emotional deficits that are characteristic of many human psychiatric disorders.
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Nathan J. Emery. (2005). The Evolution of Social Cognition. In The Cognitive Neuroscience of Social BehaviourGarten. Psychology Press.
Abstract: Although this bookis focusedon the cognitive neuroscience ofhuman social behaviour, an
understandingofsocial cognition in non-human animals is critical for unravellingthe neural basis of
social cognition in humans as well as the selective pressures that have shapedthe evolution ofcomplex
social cognition. Thanks to methodological limitations, we know little about the relationships between
certain biochemical andelectrophysiological properties ofthe human brain andhow theycompute the
behaviour andmental states ofother individuals. Traditional techniques for examiningneural function
in humans, such as event-relatedpotentials (ERP),positron emission tomography(PET),and
functional magnetic resonance imaging(fMRI),are constrainedbythe fact that subjects are placed
either into an immoveable scanner with a lot ofbackgroundnoise or wiredup with dozens of
electrodes that are sensitive to slight movements. The possibilityofscanningor recordingbrain waves
from two individuals that are physicallyinteractingsociallyis technicallyimpossible at present
(however, see Montague et al, 2002 for a new methodfor simultaneouslyscanningtwo individuals
interactingvia a computer).
The onlywayto understandthe neurocognitive architecture ofhuman social behaviour is to examine
similar social processes in both human andnon-human animal minds andmake comparisons at the
species level. An additional argument is that traditional human socio-cognitive tasks are dependent on
the use ofstories, cartoons andverbal cues andinstructions (Heberlein & Adolphs, this volume)which
themselves will elicit specific neural responses that have to be eliminatedfrom neural responses
specificallyrelatedto mindreading. Therefore, the development ofnon-verbal tasks wouldprovide a
breakthrough for studies in non-linguistic animals, pre-verbal human infants andhuman cognitive
neuroimaging.
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Napolitano, F., De Rosa, G., Braghieri, A., Grasso, F., Bordi, A., & Wemelsfelder, F. (2008). The qualitative assessment of responsiveness to environmental challenge in horses and ponies. Appl. Anim. Behav. Sci., 109(2-4), 342–354.
Abstract: The responsiveness of 10 horses and 10 ponies to environmental challenge (represented by an open field test) was assessed using a qualitative approach based on free choice profiling methodology (FCP), which gives observers complete freedom to choose their own descriptive terms. Data were analysed with generalised Procrustes analysis (GPA), a multivariate statistical technique associated with FCP. A cross-validation of the outcomes of this approach to data recorded through quantitative behaviour analysis, and through a questionnaire given to the animals' owner/riding instructor, was also performed using principal component analysis (PCA). Twelve undergraduate students generated their own descriptive vocabularies, by watching 20 horse/pony video clips lasting 2.5 min each. GPA showed that the consensus profile explained a high percentage of variation among the 12 observers, and differed significantly from the mean randomised profile (p < 0.001). Two main dimensions of the consensus profile were identified, explaining 60% and 5.2% of the variation between animals, respectively. The 12 observer word charts interpreting these dimensions were semantically consistent, as they all converged towards the same meaning, albeit using different terms. The most used term to describe the positive end of axis 1 was “quiet”, whereas “attentive” was the best positive descriptor of axis 2. The most frequently used descriptors for the negative ends of axes 1 and 2 were “nervous” and “bored”, respectively. Thus, axis 1 was labelled as “quiet/nervous” and axis 2 was named as “attentive/bored”. A marked effect of animal category was observed on the scores of the animals on the first dimension (p < 0.001). Horses received significantly higher scores, and were thus assessed as more quiet and calm, than ponies. Conversely, ponies tended to receive lower scores on the second dimension (p < 0.12), therefore they appeared less curious and attentive. The results of the PCA showed that the variables from different types of measurement clearly had meaningful relationships. For instance, the variables with the highest loading on the positive end of axis 1 were all indicative of tractable and docile animals, whereas axis 2 showed high loadings on the positive end for variables indicating attentive animals. Qualitative behaviour assessment proved to be an appropriate methodology for the study of horse behavioural responsiveness, in that it provided a multifaceted characterisation of horse behavioural expression that was in agreement with other quantitative and subjective assessments of the animals' behaviour.
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Nagy, K., Schrott, A., & Kabai, P. (2008). Possible influence of neighbours on stereotypic behaviour in horses. Appl. Anim. Behav. Sci., 111(3-4), 321–328.
Abstract: Revealing risk factors of abnormal stereotypic behaviour (ASB) in horses can help in the design of protective measures. Previous epidemiological studies indicate that social isolation, housing, management conditions, and feeding regime have a strong effect on developing ASB. The common belief that exposure to a stereotypic horse increases the risk of ASB has never been substantiated. Here we report that a generalised linear mixed models (GLMM) analysis of data on 287 horses of nine riding schools revealed that exposure to a stereotypic neighbour is a significant risk factor for performing stereotypy. Also, aggressive behaviour towards other horses increased the odds of stereotypy in the aggressor. These correspondences are unlikely to be a riding-school effect, because riding schools were treated as random factor in the GLMM. Risk factors identified by epidemiological studies cannot be treated as causal agents without independent evidence. Our aim in presenting these findings was to draw attention to the possibility of neighbour effects so that other researchers would include this variable in their surveys.
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Nagy, K., Bodó, G., Bárdos, G., Harnos, A., & Kabai, P. (2009). The effect of a feeding stress-test on the behaviour and heart rate variability of control and crib-biting horses (with or without inhibition). Appl. Anim. Behav. Sci., 121(2), 140–147.
Abstract: Crib-biting is a form of oral stereotypy affecting 4-5% of horses. Once fixed, crib-biting is difficult to eliminate by behaviour therapy, however, its performance can be inhibited by collar or surgery treatment (modified Forssell's procedure). Although surgical intervention is widespread, the effects on stress coping in horses have not been studied. In the present study we evaluated changes in behaviour response and heart rate variability in 9 control, 10 crib-biting, 10 collar and 11 surgically treated horses in a feeding stress-test, in which a feeding-bowl was placed in front but out of the reach of the horses, from which tidbits were given 3 times. We found that stress triggers high oral activity, mainly cribbing in crib-biting horses, elevates other forms of oral activities in the inhibited groups and does not affect oral activities of controls. Instead of performing oral activities, control horses tended to target an unavailable feeding-bowl by pawing or head-tossing. Changes in stress level were indistinguishable in controls and crib-biters as heart rate variability returned to baseline values in both groups. In contrast, horses inhibited to perform crib-biting showed elevated stress level throughout the test period. Our results suggest that crib-biting may develop to cope with stress, and such coping function diminishes when inhibited.
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Nagy, K., Bodó, G., Bárdos, G., Bánszky, N., & Kabai, P. (2010). Differences in temperament traits between crib-biting and control horses. Appl. Anim. Behav. Sci., 122(1), 41–47.
Abstract: Recent studies have suggested that crib-biting in horses is associated with diminished capacity of learning or coping with stress. Such findings raise the question whether trainability, which is fundamentally important in practice, could also be affected by stereotypic behaviour. Trainability of a horse is difficult to assess in simple tests, however, it is reliably estimated by experienced riders. To assess trainability and other characteristics related to that, a questionnaire survey was conducted with the owners of 50 crib-biting and 50 control horses. Where possible, control horses were selected from the same establishment as crib-biters. Groups did not differ significantly regarding age, breed, gender, training level or usage. Principal component analysis revealed three main factors which can be labelled as [`]Anxiety', [`]Affability' and [`]Trainability'. The [`]Anxiety' factor consisted of the items [`]Nervousness', [`]Excitability', [`]Panic', [`]Inconsistent emotionality', [`]Vigilance', [`]Skittishness', and [`]Timidity'. [`]Affability' consisted of [`]Friendliness toward people', [`]Cooperation', [`]Docility' and [`]Friendliness toward horses'. [`]Trainability' involved [`]Concentration', [`]Trainability', [`]Memory', and [`]Perseverance'. Temperament traits were not affected by age, gender, breed or training level, but the usage of the horse and the presence of crib-biting behaviour had significant effects. Competition horses had lower level of [`]Anxiety' (p = 0.032) and higher level of [`]Trainability' (p = 0.068) than leisure horses. Crib-biting horses had significantly lower level of [`]Anxiety' than control horses (p < 0.001), while [`]Trainability' and [`]Affability' did not differ between groups (p = 0.823 and p = 0.543, respectively). Competition horses are more often exposed to novel environment and to frightening stimuli (e.g. colourful obstacles) than leisure horses and therefore might have also become more habituated to these types of stimuli. Coping with novel situation may be enhanced by defusing nervous behaviour by the more experienced riders of competition. Previous studies indicated crib-biting horses to be less reactive when challenged as compared to control horses. We suggest that the virtual calmness and lower nervousness of the crib-biting horses might be due to the passive coping style of these animals. [`]Affability' of horses might be more related to housing and management conditions than to crib-biting. Contrary to expectations, scores on [`]Trainability' had not coincided with the impaired learning of crib-biting horses reported in laboratory tests. However, previous behavioural tests on equine learning rarely had a direct relevance to the training abilities of the horses. Our results do not support crib-biting stereotypy to affect performance in training, which is a complex learning process involving cooperation and docility in the social environment.
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