Abstract: A wide variety of maternal, social and sexual bonding strategies have been described across mammalian species, including humans. Many of the neural and hormonal mechanisms that underpin the formation and maintenance of these bonds demonstrate a considerable degree of evolutionary conservation across a representative range of these species. However, there is also a considerable degree of diversity in both the way these mechanisms are activated and in the behavioural responses that result. In the majority of small-brained mammals (including rodents), the formation of a maternal or partner preference bond requires individual recognition by olfactory cues, activation of neural mechanisms concerned with social reward by these cues and gender-specific hormonal priming for behavioural output. With the evolutionary increase of neocortex seen in monkeys and apes, there has been a corresponding increase in the complexity of social relationships and bonding strategies together with a significant redundancy in hormonal priming for motivated behaviour. Olfactory recognition and olfactory inputs to areas of the brain concerned with social reward are downregulated and recognition is based on integration of multimodal sensory cues requiring an expanded neocortex, particularly the association cortex. This emancipation from olfactory and hormonal determinants of bonding has been succeeded by the increased importance of social learning that is necessitated by living in a complex social world and, especially in humans, a world that is dominated by cultural inheritance. © 2006 The Royal Society.

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.

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.

Abstract: Abstract  Previous laboratory studies on social learning suggest that some animals can learn more readily if they first observe a conspecific demonstrator perform the task unsuccessfully and so fail to obtain a food reward than if they observe a successful demonstrator that obtains the food. This effect may indicate a difference in how easily animals are able to associate different outcomes with the conspecific or could simply be the result of having food present in only some of the demonstrations. To investigate we tested a scatter-hoarding mammal, the eastern grey squirrel, on its ability to learn to choose between two pots of food after watching a conspecific remove a nut from one of them on every trial. Squirrels that were rewarded for choosing the opposite pot to the conspecific chose correctly more frequently than squirrels rewarded for choosing the same pot (a feature-negative effect). Another group of squirrels was tested on their ability to choose between the two pots when the rewarded option was indicated by a piece of card. This time, squirrels showed no significant difference in their ability to learn to choose the same or the opposite pot. The results add to anecdotal reports that grey squirrels can learn by observing a conspecific and suggest that even when all subjects are provided with demonstrations with the same content, not all learning occurs equally. Prior experience or expectations of the association between a cue (a conspecific) and food influences what can be learned through observation whilst previously unfamiliar cues (the card) can be associated more readily with any outcome.

Abstract: Comparative experimental studies of imitative learning have focused mainly on primates and birds. However, cetaceans are promising candidates to display imitative learning as they have evolved in socioecological settings that have selected for large brains, complex sociality, and coordinated predatory tactics. Here we tested imitative learning in killer whales, Orcinus orca. We used a ‘do-as-other-does’ paradigm in which 3 subjects witnessed a conspecific demonstrator’s performance that included 15 familiar and 4 novel behaviours. The three subjects (1) learned the copy command signal ‘Do that’ very quickly, that is, 20 trials on average; (2) copied 100 % of the demonstrator’s familiar and novel actions; (3) achieved full matches in the first attempt for 8–13 familiar behaviours (out of 15) and for the 2 novel behaviours (out of 2) in one subject; and (4) took no longer than 8 trials to accurately copy any familiar behaviour, and no longer than 16 trials to copy any novel behaviour. This study provides experimental evidence for body imitation, including production imitation, in killer whales that is comparable to that observed in dolphins tested under similar conditions. These findings suggest that imitative learning may underpin some of the group-specific traditions reported in killer whales in the field.