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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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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.
Keywords: Animal welfare; Emotion; Separation; Social attachment; Weaning
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Newton-Fisher, N. E., & Lee, P. C. (2011). Grooming reciprocity in wild male chimpanzees. Anim. Behav., 81(2), 439–446.
Abstract: Understanding cooperation between unrelated individuals remains a central problem in animal behaviour; evolutionary mechanisms are debated, and the importance of reciprocity has been questioned. Biological market theory makes specific predictions about the occurrence of reciprocity in social groups; applied to the social grooming of mammals, it predicts reciprocity in the absence of other benefits for which grooming can be exchanged. Considerable effort has been made to test this grooming trade model in nonhuman primates; such studies show mixed results, but may be confounded by kin effects. We examined patterns of reciprocity within and across bouts, and tested predictions of the grooming trade model, among wild male chimpanzees, Pan troglodytes: a system with negligible kin effects. In accord with the model's expectations, we found that some grooming was directed by lower- to higher-ranked individuals, and that, on average, higher-ranked individuals groomed more reciprocally. We found no support, however, for a prediction that more reciprocity should occur between individuals close in rank. For most dyads, reciprocity of effort occurred through unbalanced participation in grooming bouts, but reciprocity varied considerably between dyads and only a small proportion showed strongly reciprocal grooming. Despite this, each male had at least one reciprocal grooming relationship. In bouts where both individuals groomed, effort was matched through mutual grooming, not alternating roles. Our results provide mixed support for the current grooming trade, biological market model, and suggest that it needs to incorporate risks of currency inflation and cheating for species where reciprocity can be achieved through repeated dyadic interactions.
Keywords: altruism; biological market theory; Budongo; chimpanzee; grooming; Pan troglodytes
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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.
Keywords: Learning; Horse; Equine; Discrimination; Training
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Nicol, C. J. (2006). How animals learn from each other. Appl. Anim. Behav. Sci., 100(1-2), 58–63.
Abstract: This paper explores ways by which animals may learn from one another, using examples drawn mostly from the chicken, an animal for which social learning is likely to be less dangerous than individual learning. In early life, the behaviour of the hen is important in encouraging chicks to peck at edible items. Maternal display not only attracts chicks to profitable food items, but also redirects their attention away from harmful or non-profitable items. Older chicks can enhance their foraging success by observing the behaviour of conspecifics within their own social group. Hens have been trained to perform a novel behaviour (key-pecking for food) by observation of a trained demonstrator bird. Moreover, observers learnt most from watching dominant demonstrators. Thus the ability to learn from others is not `fixed', but depends on the context and the social identity of both the observer and the demonstrator.
Keywords: Social learning; Chickens; Demonstrators; Dominance
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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., & Badnell-Waters, A. J. (2005). Suckling behaviour in domestic foals and the development of abnormal oral behaviour. Anim. Behav., 70(1), 21–29.
Abstract: We investigated how the behaviour of domestic foals, Equus caballus, living at pasture with their dams was associated with foal gender, mare rank and the development of abnormal oral behaviour, both during the preweaning period, and over a period of up to 4 years postweaning. A population of 186 foals belonging to private owners and commercial studs was studied. The behaviour of male and female foals hardly differed, but mare rank affected patterns of foal social interaction and suckling behaviour, with foals of subordinate mares involved in more affiliative interactions. These foals also spent more time in perisuckling activities such as teat nuzzling than foals of other mares. During the study, 18 foals developed abnormal oral behaviour before weaning and 42 foals developed abnormal oral behaviour after weaning. The development of abnormal oral behaviour was associated with suckling behaviour in a variety of ways. Foals that had already developed abnormal oral behaviour at the time of the preweaning observations were involved in more suckling terminations within bouts than normal foals or foals that developed future abnormal behaviour, and pushing the udder with the muzzle was most frequent in these foals. Foals that had no current abnormal oral behaviour, but that would develop this in the future, spent more time suckling and twice as much time teat nuzzling as other foals. The results add to the growing evidence of associations between digestive function and abnormal oral behaviour in horses.
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Nicol, C. J., Badnell-Waters, A. J., Bice, R., Kelland, A., Wilson, A. D., & Harris, P. A. (2005). The effects of diet and weaning method on the behaviour of young horses. Appl. Anim. Behav. Sci., 95(3-4), 205–221.
Abstract: The effects of diet on horse behaviour have not previously been quantified in detail. In this study, we examined the behaviour of 17 foals from the age of 2 to 40 weeks. Each foal received either a starch and sugar (SS) diet or a fat and fibre (FF) diet. The two diets contained similar digestible energy, crude protein and micronutrients, but differed in the fat and non-structural carbohydrate balance. The baseline behaviour of the foals was observed every 2 weeks by focal animal sampling. Additional behavioural observations were conducted when the foals were weaned by one of two methods. Approximately 2 months after weaning, the temperament and tractability of the young horses was assessed using standardised tests. Responses to a novel object, to a novel person, and during a handling test were observed and quantified. Horses grew well on both diets with no apparent effects of diet on growth rate or baseline behaviour. Immediately after weaning, horses receiving the FF diet cantered less frequently (F = 5.10; p < 0.05), for a shorter duration (F = 7.23; p < 0.05) and appeared to be more settled. Foals that were barn-weaned appeared more stressed than foals that were paddock-weaned. In the temperament tests, horses receiving the FF diet spent significantly more time investigating (F = 6.78; p < 0.05), and less time looking at (F = 7.93; p < 0.05), the novel object than horses receiving the SS diet. They also spent less time walking away from the novel person (F = 5.16; p < 0.05) and their time taken to complete the handling test was significantly lower (F = 8.72; p = 0.01). Overall, the horses that received the FF diet appeared less distressed immediately after weaning and seemed calmer and more inquisitive during a range of temperament tests.
Keywords: Horse; Diet; Weaning; Temperament test
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Nicol, C. J., & Pope, S. J. (1999). The effects of demonstrator social status and prior foraging success on social learning in laying hens. Anim. Behav., 57(1), 163–171.
Abstract: Opportunities for social learning within a group of animals are likely to be influenced by the social dynamics of that group. Some individuals may be more influential demonstrators than others even when there are no differences in their skill level or performance. In this study of domestic hens,Gallus gallus domesticus, differences in demonstrator salience were examined. From 24 separate flocks we selected as demonstrators a dominant cockerel, a dominant hen, a mid-ranking hen or a subordinate hen. Demonstrators were pretrained to perform an operant discrimination task to obtain food. Six observers from each flock individually watched the demonstrator perform the task for four 5-min sessions held on consecutive days. On the fifth day observers were tested individually in the operant chamber. We analysed data from 19 flocks, where there were no quantitative differences in demonstrator performance. Observer hens of relatively high social status performed more correct operant pecks than observer hens of relatively low social status. Demonstrator category also had a significant effect on subsequent observer behaviour. Hens that had observed cockerels performed very few general pecks or operant pecks. Hens that had observed dominant hens performed more operant pecks, but hens that had observed sub-ordinate hens performed more general pecks in the chamber. The results suggested either that there was an interaction between dominance and gender in demonstrator salience or that dominant hens might have been influential because of some factor imperfectly associated with their dominance status. A possible candidate was the foraging ability of the dominant hens. In a second experiment using the same protocol, we manipulated the prior foraging success of dominant hens from four additional flocks but this had no significant effect on their subsequent influence as demonstrators.
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NICOLA J. ROONEY & JOHN W. S. BRADSHAW. (2005). Social cognition in the domestic dog: behaviour of spectators towards participants in interspecific games. Anim. Behav., 72(2), 343–352.
Abstract: previous termSocial cognition,next term in particular the derivation of previous termsocialnext term information from observation of interactions between members of a previous termsocialnext term group, has been widely investigated in primates, but it has received little attention in other previous termsocialnext term mammals, although it has been anecdotally reported in the previous termdomestic dog,next term Canis familiaris. We recorded the behaviour of previous termdogsnext term (“spectators”) that had observed controlled interactions between a human and a previous termdognext term (the “demonstrator”) competing for an object, and that were subsequently allowed to interact freely with both participants. When the competitions were playful, as indicated by signals performed by the human, the spectator was more likely to approach the winner first and/or more rapidly, suggesting that winners of games are perceived as desirable previous termsocialnext term partners. When the human did not perform play signals, changing the previous termsocialnext term context from play to contest over a resource, spectators were slower to approach either of the participants, suggesting that participants in contests were less desirable as previous termsocialnext term partners than participants in games. If the previous termdognext term was prevented from seeing the game, it still reacted differently to the winner and the loser, but its behaviour was not the same as after games that it had seen. We conclude that spectator previous termdogsnext term gain information from the players' subsequent behaviour as well as from direct observation of the game.
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