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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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Merola, I., Lazzaroni, M., Marshall-Pescini, S., & Prato-Previde, E. (2015). Social referencing and cat–human communication. Anim. Cogn., 18(3), 639–648.
Abstract: Cats’ (Felis catus) communicative behaviour towards humans was explored using a social referencing paradigm in the presence of a potentially frightening object. One group of cats observed their owner delivering a positive emotional message, whereas another group received a negative emotional message. The aim was to evaluate whether cats use the emotional information provided by their owners about a novel/unfamiliar object to guide their own behaviour towards it. We assessed the presence of social referencing, in terms of referential looking towards the owner (defined as looking to the owner immediately before or after looking at the object), the behavioural regulation based on the owner’s emotional (positive vs negative) message (vocal and facial), and the observational conditioning following the owner’s actions towards the object. Most cats (79 %) exhibited referential looking between the owner and the object, and also to some extent changed their behaviour in line with the emotional message given by the owner. Results are discussed in relation to social referencing in other species (dogs in particular) and cats’ social organization and domestication history.
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McVey, A., Wilkinson, A., & Mills, D. S. (2018). Social learning in horses: the effect of using a group leader demonstrator on the performance of familiar conspecifics in a detour task. Applied Animal Behaviour Science, .
Abstract: Learning through the observation of others allows the transfer of information without the costs incurred during individual trial and error learning. Horses (Equus caballus) are a highly social species, which might be expected to be capable of learning from others, but experimental findings are inconsistent, and potentially confounded by social facilitation effects not related directly to the learning of the task. We refined the methods used in previous equine social learning studies, to examine and distinguish specific social influences on learning of a task: we used predefined group leaders rather than agonistically dominant individuals to demonstrate a detour task to familiar conspecific observers; in addition we had two control groups: a non-observer (true control) and a group with the demonstrator simply present at the goal (social facilitation control). 44 socially kept horses were allocated to one of the three test conditions and took part in five trials each. Success rate, latency and detour direction were recorded. There was no significant difference between the three groups in the likelihood of them succeeding in the task nor latency to succeed; however there was a significant difference in the route chosen by the groups, with the true control choosing the side with the entrance gate significantly more than either the observer group or social facilitation group. Both of the latter two groups chose to go in the same direction relative to themselves, regardless of which side the gate was. Seven out of nine horses in the observer group chose the same direction as their demonstrator every time. Our results show a significant role of social facilitation on detour behaviour and highlight the importance of including adequate controls for simpler cognitive influences on behaviour before claims can be made about the specific learning of motor actions or goal directed behaviour. Social cues may be important to horses if the task is sufficiently challenging and motivationally important, so future work should consider more demanding, but ecologically relevant situations, in order to maximise the potential revelation of social learning effects which do not depend on simple local or stimulus enhancement effects.
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Lefebvre, L. (1995). Ecological correlates of social learning: problems and solutions for the comparative method. Behav. Process., 35(1-3), 163–171.
Abstract: Interspecific variation in learning and cognition is often accounted for by adaptive specialization, an ecological framework where variation between species in the environmental problems they face is thought to select for quantitatively and/or qualitatively different abilities. Adaptive specialization theory relies on the comparative method for testing its hypotheses and assumes a naturally selected basis for the predicted differences. This review examines social learning as a specialization to group-living and scramble feeding competition. It points out one important problem with current studies in the area, the lack of quantitative controls for confounding variables that may cause type 1 or 2 error in comparative tests. A linear regression technique is proposed to measure and remove interspecific differences on control tests for which there is no predicted adaptive specialization; as in other areas of comparative biology, the adaptive prediction is then made on the residual deviation from the regression of these confounding variables. Examples are given from research on opportunistic Columbids, the group-living feral pigeon Columbia livia, and the territorial Zenaida dove, Zenaida aurita.
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Leadbeater, E. (2015). What evolves in the evolution of social learning? J Zool, 295(1), 4–11.
Abstract: Social learning is fundamental to social life across the animal kingdom, but we still know little about how natural selection has shaped social learning abilities on a proximate level. Sometimes, complex social learning phenomena can be entirely explained by Pavlovian processes that have little to do with the evolution of sociality. This implies that the ability to learn socially could be an exaptation, not an adaptation, to social life but not that social learning abilities have been left untouched by natural selection. I discuss new empirical evidence for associative learning in social information use, explain how natural selection might facilitate the associative learning process and discuss why such studies are changing the way that we think about social learning.
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Krueger., K., & Farmer, K. (2018). Social learning in Horses: Differs from individual learning only in the learning stimulus and not in the learning mechanisms. In 14th Meeting of the Internatinoal Society for Equitation Science.
Abstract: Equine welfare can be enhanced by applying species specific training. This may incorporate social learning, as horses are highly social and social stimuli are of primary importance. Social learning is comparable to individual learning in its learning mechanisms, differing primarily in the way it is stimulated. Our initial study showed that horses of different breeds (N = 38) follow humans after observing other horses doing so, but only if the observed horse was familiar to and higher ranking than the observer (Fisher's exact test: N = 12, P = 0.003). A second study showed that horses and ponies (N = 25) learned to pull a rope to open a feeding apparatus after observing demonstrations by conspecifics, again, only if the demonstrating horse was older and higher ranking than the observer (Fisher's combination test, N = 3, v2 = 27.71, p = 0.006). Our third approach showed that horses and ponies (N = 24) learned to press a switch to open a feeding apparatus after observing a familiar person (GzLM: N = 24, z = 2.33, P = 0.02). Most recently, we confronted horses and ponies (N = 50) with persons demonstrating different techniques for opening a feeding apparatus. In this study we investigated whether the horses would copy the demonstrators' techniques or apply their own. Here only some horses copied the technique, and most of the successful learners used their mouths irrespective of the demonstrators' postures (Chi Square Test: N = 40, df = 2, χ2 = 31.4, p < 0.001). In all the approaches social stimuli elicited learning processes in the test horses, while only a few individuals in the control groups mastered the tasks by individual learning. The following behaviour observed in the initial study may have been facilitated by a social stimuli (social facilitation), and the opening of the feed boxes in the subsequent studies appear to be mostly the result of enhancement (social enhancement). Some horses may have used the social stimuli at first and continued their learning process by individual trial and error. However, the horses were also selective in whom and some in how to copy. This may have been conditioned (socially conditioned) or the result of simple forms of reasoning on the reliability of the particular information provided by demonstrators of certain social ranks or social positions, as high ranking and familiar horses and familiar persons were copied and some imitated exactly.
Lay person message: Traditional riding instructions suggest that horses learn by observing other horses. For example, older, more experienced driving horses are used for initial training of young driving horses. We have shown that horses indeed use learning stimuli provided by other horse, as well as by humans. Horses readily accept stimuli observed in high ranking and familiar horses, and familiar persons. Such stimuli elicit learning processes which are comparable to individual learning. We suggest applying social learning whenever possible, as it is much faster and less stressful than individual learning, where learners experience negative outcomes in trial and error learning.
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Krueger, K., Farmer, K., & Heinze, J. (2014). The effects of age, rank and neophobia on social learning in horses. Anim. Cogn., 17(3), 645–655.
Abstract: Social learning is said to meet the demands of complex environments in which individuals compete over resources and co-operate to share resources. Horses (Equus caballus) were thought to lack social learning skills because they feed on homogenously distributed resources with few reasons for conflict. However, the horse’s social environment is complex, which raises the possibility that its capacity for social transfer of feeding behaviour has been underestimated. We conducted a social learning experiment using 30 socially kept horses of different ages. Five horses, one from each group, were chosen as demonstrators, and the remaining 25 horses were designated observers. Observers from each group were allowed to watch their group demonstrator opening a feeding apparatus. We found that young, low ranking, and more exploratory horses learned by observing older members of their own group, and the older the horse, the more slowly it appeared to learn. Social learning may be an adaptive specialisation to the social environment. Older animals may avoid the potential costs of acquiring complex and potentially disadvantageous feeding behaviours from younger group members. We argue that horses show social learning in the context of their social ecology, and that research procedures must take such contexts into account. Misconceptions about the horse’s sociality may have hampered earlier studies.
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Krueger, K. (2015). Social learning and innovative behaviour in horses. In Proceedings of the 3. International Equine Science Meeting. Wald: Xenophon Publishing.
Abstract: The evaluation of important parameters for measuring the horses’ cognitive capacities is one of the central topics of the equine behaviour team at Nürtingen-Geislingen University. Social complexity has been said to be one of the settings in which needs for cognitive capacities arise in animals. A variety of studies throughout the last two decades proved the horses’ social complexity to be far more elaborate than previously assumed. Horses form social bonds for the protection of offspring, intervene in encounters of others, identify group mates individually and easily orientate in a fission fusion society.
In such socially complex societies, animals will benefit from learning socially. In many bird and primate species the degree of social complexity correlates nicely with the species abilities for social learning. Social learning was, therefore, argued to be an indicator for elaborate mental capacities in animals. We were delighted to prove that horses actually copy social behaviour and techniques for operating a feeding apparatus from older and higher ranking group members. In a recent study we found young horses, at the age of 3 to 12, to copy the operation of a feeding apparatus from a human demonstrator. Social learning seems to work nicely in horses when the social background of the animals is considered.
The degree to which individual animals adapt to changes in their social or physical environment by finding innovative solution appears to be the other side of the coin, of whether animals adjust to challenges by social learning. It is not very astonishing, that along with the animals’ social complexity and their ability to learn socially also the degree to which they show innovative behaviour was claimed to be one of the most important demonstrations of advanced cognitive capacities. In a recent approach, we started to ask horse owners and horse keepers in many countries to tell us about unusual behaviour of their horses via a web site (http://innovative-behaviour.org). To date, we received 204 cases of innovative behaviour descriptions from which six cases were clear examples of tool use or borderline tool use. We categorized the innovative behaviours into the classes, a) innovations to gain food, b) innovations to gain freedom, c) social innovations, d) innovations to increase maintenance, and e) innovations that could not be clearly assigned to a category. About 20% of the innovative horses showed more than one innovation. These animals could be termed “true innovators”. Again, young horses were more innovative than older ones with the age group 5 – 9 showing the highest number of innovative behaviour descriptions.
In a nutshell, the horses’ cognitive capacities appear to be underestimated throughout the last decades. The horses’ social complexity is far more elaborate than previously assumed, horses learn socially from conspecific and humans, some of them demonstrate innovative behaviour adaptations to their environment and even simple forms of tool use.
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Hopewell, L., Leaver, L., Lea, S., & Wills, A. (2010). Grey squirrels (Sciurus carolinensis) show a feature-negative effect specific to social learning. Anim. Cogn., 13(2), 219–227.
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.
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Hasenjager, M. J., & Dugatkin, L. A. Social Network Analysis in Behavioral Ecology. Advances in the Study of Behavior. Academic Press.
Abstract: Abstract In recent years, behavioral ecologists have embraced social network analysis (SNA) in order to explore the structure of animal societies and the functional consequences of that structure. We provide a conceptual introduction to the field that focuses on historical developments, as well as on novel insights generated by recent work. First, we discuss major advances in the analysis of nonhuman societies, culminating in the use of SNA by behavioral ecologists. Next, we discuss how network-based approaches have enhanced our understanding of social structure and behavior over the past decade, focusing on: (1) information transmission, (2) collective behaviors, (3) animal personality, and (4) cooperation. These behaviors and phenomena possess several features—e.g., indirect effects, emergent properties—that network analysis is well equipped to handle. Finally, we highlight recent developments in SNA that are allowing behavioral ecologists to address increasingly sophisticated questions regarding the structure and function of animal sociality.
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