What imitation tells us about social cognition: a rapprochement between developmental psychology and cognitive neuroscience

Both developmental and neurophysiological research suggest a common coding between perceived and generated actions. This shared representational network is innately wired in humans. We review psychological evidence concerning the imitative behaviour of newborn human infants. We suggest that the mechanisms involved in infant imitation provide the foundation for understanding that others are 'like me' and underlie the development of theory of mind and empathy for others. We also analyse functional neuroimaging studies that explore the neurophysiological substrate of imitation in adults. We marshal evidence that imitation recruits not only shared neural representations between the self and the other but also cortical regions in the parietal cortex that are crucial for distinguishing between the perspective of self and other. Imitation is doubly revealing: it is used by infants to learn about adults, and by scientists to understand the organization and functioning of the brain.     

Copying You Copying Me: Interpersonal Motor Co-Ordination Influences Automatic Imitation

Moving in a co-ordinated fashion with another individual changes our behaviour towards them; we tend to like them more, find them more attractive, and are more willing to co-operate with them. It is generally assumed that this effect on behaviour results from alterations in representations of self and others. Specifically, through neurophysiological perception-action matching mechanisms, interpersonal motor co-ordination (IMC) is believed to forge a neural coupling between actor and observer, which serves to blur boundaries in conceptual self-other representations and causes positive views of the self to be projected onto others. An investigation into this potential neural mechanism is lacking, however. Moreover, the specific components of IMC that might influence this mechanism have not yet been specified. In the present study we exploited a robust behavioural phenomenon – automatic imitation – to assess the degree to which IMC influences neural action observation-execution matching mechanisms. This revealed that automatic imitation is reduced when the actions of another individual are perceived to be synchronised in time, but are spatially incongruent, with our own. We interpret our findings as evidence that IMC does indeed exert an effect on neural perception-action matching mechanisms, but this serves to promote better self-other distinction. Our findings demonstrate that further investigation is required to understand the complex relationship between neural perception-action coupling, conceptual self-other representations, and social behaviour.     

Self–other control processes in social cognition: from imitation to empathy

We review the evidence that an ability to achieve a precise balance between representing the self and representing other people is crucial in social interaction. This ability is required for imitation, perspective-taking, theory of mind and empathy; and disruption to this ability may contribute to the symptoms of clinical and sub-clinical conditions, including autism spectrum disorder and mirror-touch synaesthesia. Moving beyond correlational approaches, a recent intervention study demonstrated that training participants to control representations of the self and others improves their ability to control imitative behaviour, and to take another''s visual perspective. However, it is unclear whether these effects apply to other areas of social interaction, such as the ability to empathize with others. We report original data showing that participants trained to increase self–other control in the motor domain demonstrated increased empathic corticospinal responses (Experiment 1) and self-reported empathy (Experiment 2), as well as an increased ability to control imitation. These results suggest that the ability to control self and other representations contributes to empathy as well as to other types of social interaction.     

Homo imitans? Seven reasons why imitation couldn't possibly be associative

Many comparative and developmental psychologists believe that we are Homo imitans; humans are more skilled and prolific imitators than other animals, because we have a special, inborn ‘intermodal matching’ mechanism that integrates representations of others with representations of the self. In contrast, the associative sequence learning (ASL) model suggests that human infants learn to imitate using mechanisms that they share with other animals, and the rich resources provided by their sociocultural environments. This article answers seven objections to the ASL model: (i) it presents evidence that newborns do not imitate; (ii) argues that infants receive a plentiful supply of the kind of experience necessary for learning to imitate; (iii) suggests that neither infants nor adults can imitate elementally novel actions; (iv) explains why non-human animals have a limited capacity for imitation; (v) discusses the goal-directedness of imitation; (vi) presents evidence that improvement in imitation depends on visual feedback; and (vii) reflects on the view that associative theories steal ‘the soul of imitation’. The empirical success of the ASL model indicates that the mechanisms which make imitation possible, by aligning representations of self with representations of others, have been tweaked by cultural evolution, not built from scratch by genetic evolution.     

Evolving intentions for social interaction: from entrainment to joint action

This article discusses four different scenarios to specify increasingly complex mechanisms that enable increasingly flexible social interactions. The key dimension on which these mechanisms differ is the extent to which organisms are able to process other organisms' intentions and to keep them apart from their own. Drawing on findings from ecological psychology, scenario 1 focuses on entrainment and simultaneous affordance in 'intentionally blind' individuals. Scenario 2 discusses how an interface between perception and action allows observers to simulate intentional action in others. Scenario 3 is concerned with shared perceptions, arising through joint attention and the ability to distinguish between self and other. Scenario 4 illustrates how people could form intentions to act together while simultaneously distinguishing between their own and the other's part of a joint action. The final part focuses on how combining the functionality of the four mechanisms can explain different forms of social interactions. It is proposed that basic interpersonal processes are put to service by more advanced functions that support the type of intentionality required to engage in joint action, cultural learning, and communication.     

Grist and mills: on the cultural origins of cultural learning

Cumulative cultural evolution is what 'makes us odd'; our capacity to learn facts and techniques from others, and to refine them over generations, plays a major role in making human minds and lives radically different from those of other animals. In this article, I discuss cognitive processes that are known collectively as 'cultural learning' because they enable cumulative cultural evolution. These cognitive processes include reading, social learning, imitation, teaching, social motivation and theory of mind. Taking the first of these three types of cultural learning as examples, I ask whether and to what extent these cognitive processes have been adapted genetically or culturally to enable cumulative cultural evolution. I find that recent empirical work in comparative psychology, developmental psychology and cognitive neuroscience provides surprisingly little evidence of genetic adaptation, and ample evidence of cultural adaptation. This raises the possibility that it is not only 'grist' but also 'mills' that are culturally inherited; through social interaction in the course of development, we not only acquire facts about the world and how to deal with it (grist), we also build the cognitive processes that make 'fact inheritance' possible (mills).     

Neural basis of self and other representation in autism: an FMRI study of self-face recognition

Background

Autism is a developmental disorder characterized by decreased interest and engagement in social interactions and by enhanced self-focus. While previous theoretical approaches to understanding autism have emphasized social impairments and altered interpersonal interactions, there is a recent shift towards understanding the nature of the representation of the self in individuals with autism spectrum disorders (ASD). Still, the neural mechanisms subserving self-representations in ASD are relatively unexplored.

Methodology/Principal Findings

We used event-related fMRI to investigate brain responsiveness to images of the subjects'' own face and to faces of others. Children with ASD and typically developing (TD) children viewed randomly presented digital morphs between their own face and a gender-matched other face, and made “self/other” judgments. Both groups of children activated a right premotor/prefrontal system when identifying images containing a greater percentage of the self face. However, while TD children showed activation of this system during both self- and other-processing, children with ASD only recruited this system while viewing images containing mostly their own face.

Conclusions/Significance

This functional dissociation between the representation of self versus others points to a potential neural substrate for the characteristic self-focus and decreased social understanding exhibited by these individuals, and suggests that individuals with ASD lack the shared neural representations for self and others that TD children and adults possess and may use to understand others.     

Bodily selves in relation: embodied simulation as second-person perspective on intersubjectivity

This article addresses basic aspects of social cognition focusing on the pivotal role played by the lived body in the constitution of our experience of others. It is suggested that before studying intersubjectivity we should better qualify the notion of the self. A minimal notion of the self, the bodily self, defined in terms of its motor potentialities, is proposed. The discovery of mirror mechanisms for action, emotions and sensations led to the proposal of an embodied approach to intersubjectivity—embodied simulation (ES) theory. ES and the related notion of neural reuse provide a new empirically based perspective on intersubjectivity, viewed first and foremost as intercorporeality. ES challenges the notion that folk psychology is the sole account of interpersonal understanding. ES is discussed within a second-person perspective on mindreading.     

Impaired mirror-image imitation in Asperger and high-functioning autistic subjects

Imitation is crucial for proper development of social and communicative skills. Here, we argue that, based on an error analysis of a behavioral imitation task, adult Asperger and high-functioning autistic subjects suffer from an intriguing deficit of imitation: they lack the natural preference for imitation in a mirror-image fashion. The imitation task consisted of a simple movement sequence of putting a pen with the left or right hand into a green or a blue cup using one of two possible grips. The subjects were asked to imitate the experimenter's hand movements either using the crossed hand (e.g., the subject's right hand corresponding to the experimenter's right hand) for imitation or to imitate as if looking in a mirror (e.g., the subject's left hand corresponding to the experimenter's right hand). When people normally view other persons face-to-face, they prefer to imitate as in a mirror, and observation of mirror-image-like movements speeds up performance in nonimitative tasks. However, our autistic subjects, defective in social cognition, did not profit from mirror-image movements of others. These results provide a new insight into the difficulties that autistic subjects face in viewing and understanding actions of others.     

Representation of others' action by neurons in monkey medial frontal cortex

Successful social interaction depends on not only the ability to identify with others but also the ability to distinguish between aspects of self and others. Although there is considerable knowledge of a shared neural substrate between self-action and others' action, it remains unknown where and how in the brain the action of others is uniquely represented. Exploring such agent-specific neural codes is important because one's action and intention can differ between individuals. Moreover, the assignment of social agency breaks down in a range of mental disorders. Here, using two monkeys monitoring each other's action for adaptive behavioral planning, we show that the medial frontal cortex (MFC) contains a group of neurons that selectively encode others' action. These neurons, observed in both dominant and submissive monkeys, were significantly more prevalent in the dorsomedial convexity region of the MFC including the pre-supplementary motor area than in the cingulate sulcus region of the MFC including the rostral cingulate motor area. Further tests revealed that the difference in neuronal activity was not due to gaze direction or muscular activity. We suggest that the MFC is involved in self-other differentiation in the domain of motor action and provides a fundamental neural signal for social learning.     

To Know One's Place: Belonging and Differentiation in Alice Springs Town

Based on research in the town of Alice Springs in Central Australia the article explores how social and material aspects of the town generate meaningful understandings of oneself and differentiated others. Drawing on anthropology of place and space and analytical notions of belonging, I explore shared and divergent ways in which a range of people in town come to ‘know their place’ both in a socio-cultural sense and in the sense of relating to the built environment. Paying just as detailed attention to non-indigenous experience as scholars have long paid to indigenous lives, the article suggests that a focus on how people form material and social attachments to place can facilitate more open-ended understandings of changing forms of indigenous-settler relations than the more common focus on difference and division between categories of indigenous and non-indigenous people and domains of life.     

The ethological analysis of imitation

Theorists and experimental researchers have long debated whether animals are able to imitate. A variety of definitions of imitation have been proposed to describe this complex form of social learning. Experimental research on imitation has often been hampered by either a too loose 'anthropomorphic' approach or by too narrow 'behaviourist' definitions. At present neither associative nor cognitive theories are able to offer an exhaustive explanation of imitation in animals. An ethological approach to imitation offers a different perspective. By integrating questions on function, mechanism, development and evolution one can identify possible directions for future research. At present, however, we are still far from developing a comprehensive theory of imitation. A functional approach to imitation shows that, despite some evidence for imitative learning in food processing in apes, such learning has not been shown to be involved in the social transmission of either tool-use skills or communicative signals. Recently developed procedures offer possible ways of clarifying the role of imitation in tool use and visual communication. The role of imitation in explorative play in apes is also investigated and the available data suggest that copying during play might represent a behavioural homologue of human imitation. It is proposed that the ability to copy the behaviour of a companion is under a strong genetic influence in many social species. Many important factors have not been examined experimentally, e.g. the effect of the demonstrator, the influence of attention and memory and the ability to generalize. The potential importance of reinforcement raises the possibility that copying abilities serving divergent functions might be partly under the control of different mechanisms.     

The role of metacognition in human social interactions

Metacognition concerns the processes by which we monitor and control our own cognitive processes. It can also be applied to others, in which case it is known as mentalizing. Both kinds of metacognition have implicit and explicit forms, where implicit means automatic and without awareness. Implicit metacognition enables us to adopt a we-mode, through which we automatically take account of the knowledge and intentions of others. Adoption of this mode enhances joint action. Explicit metacognition enables us to reflect on and justify our behaviour to others. However, access to the underlying processes is very limited for both self and others and our reports on our own and others' intentions can be very inaccurate. On the other hand, recent experiments have shown that, through discussions of our perceptual experiences with others, we can detect sensory signals more accurately, even in the absence of objective feedback. Through our willingness to discuss with others the reasons for our actions and perceptions, we overcome our lack of direct access to the underlying cognitive processes. This creates the potential for us to build more accurate accounts of the world and of ourselves. I suggest, therefore, that explicit metacognition is a uniquely human ability that has evolved through its enhancement of collaborative decision-making.     

Affect,creativity, and community‐making in a Congolese song‐dance performance: or how to follow the movement of the social

In rural Punu society, song‐dance performances of rejoicing are deeply connected with forces of regeneration; they not only celebrate the sexual encounter but also, in their constituent dynamic, aim progressively to awaken a shared feeling of joy that leads to the intensification of the dancing and to the creation and diffusion of new songs. I seek to account for this regenerative potential by pursuing Turner's processual view of ritual performance, which acknowledges moments of structure and lack of structure. In this regard, I develop the dynamic of community‐making and creative flow in the performances with reference to Collins's reading of Durkheim's collective effervescence and Tarde's idea of imitation. As a whole, I argue that paying attention to rhythm and affect, inasmuch as they are a lived realization of structure and flow, is essential for understanding the revitalizing articulation between these terms and for following the ongoing movement of the social.     

What is love? The complex relation between values and practice in Vanuatu

The past decade has seen a renewed anthropological interest in values, morality, and ethics. This article engages with this field by demonstrating how values can be strategies as well as ideals, prone to destabilize social order and divide people precisely because they are thought to be shared. The concept of ‘love’, referring to everyday practices of concern and care for others, is a core value for living on Ahamb Island in Vanuatu. However, adherence to the same core value does not necessarily create an ordered social world. Analysing three ethnographic cases, one of them a dispute with fatal consequences, I propose a model for studying values that accommodates ambiguity by uniting the notion of shared social values with individual experience and strategy. A methodological argument is that it is crucial for anthropological studies of values to assess the context for people's shifting interpretations and articulations of value in practice.     

‘A good neighbour is not one that gives’: detachment,ethics, and the relational self in Kenya

Since Mauss, anthropologists have been drawn to gifts because they are understood to index the moral, often in contrast to the amorality of the market. Giving is considered to bring people closer together, weaving them into the fabric of sociality. However, in Korogocho, a low‐income informal settlement in Nairobi, women recognize the dangers of giving and of closeness. Not only could others be a burden on them, but they could also be a burden on others. Thus, women seek ways in which they can detach from one another. Beginning with the assumption of the relational self, I argue that this detachment is an ethical practice that can be best understood as what I term a ‘care for relationships’. In making this argument, this article aims to contribute to debates within the anthropology of ethics and morality concerning social relationships by showing the way in which relationships might be the explicit object of ethical practices.     

Evolving institutions for collective action by selective imitation and self-interested design

Human behavior and collective actions are strongly affected by social institutions. A question of great theoretical and practical importance is how successful social institutions get established and spread across groups and societies. Here, using institutionalized punishment in small-scale societies as an example, we contrast two prominent mechanisms - selective imitation and self-interested design - with respect to their ability to converge to cooperative social institutions. While selective imitation has received a great deal of attention in studies of social and cultural evolution, the theoretical toolbox for studying self-interested design is limited. Recently Perry, Shrestha, Vose, and Gavrilets (2018) expanded this toolbox by introducing a novel approach, which they called foresight, generalizing standard myopic best response for the case of individuals with a bounded ability to anticipate actions of their group-mates and care about future payoffs. Here we apply this approach to two general types of collective action – “us vs. nature” and “us vs. them” games. We consider groups composed by a number of regular members producing collective good and a leader monitoring and punishing free-riders. Our results show that foresight increases leaders' willingness to punish free-riders. This, in turn, leads to increased production and the emergence of an effective institution for collective action. We also observed that largely similar outcomes can be achieved by selective imitation, as argued earlier. Selective imitation by leaders (i.e. cultural group selection) outperforms self-interested design if leaders strongly discount the future. Foresight and selective imitation can interact synergistically leading to a faster convergence to an equilibrium. Our approach is applicable to many other types of social institutions and collective action.     

Spatial Group Structure as Potential Mechanism to Maintain Cooperation in Fish Shoals of Unrelated Individuals

In public goods games, group members share the benefits created through individual investments. If benefits are shared equally, individual contributions readily become altruistic, and hence, a social dilemma arises in which group interests and individual interests oppose each other. However, contributions to public goods can be self‐serving if each investor gains a disproportionate benefit from its own contribution. This scenario may hold for our study system, the interactions of shoaling‐unrelated scalefin anthias Pseudanthias squamipinnis and the ectoparasitic blenny Plagiotremus tapeinosoma. The blenny bites off pieces of skin, mucus and scales from anthias that in return may chase the blenny. Chasing the blenny represents a public good as it makes the parasite change victim species for its next attack. Laboratory experiments using artificial Plexiglas hosts suggest that one reason why individuals contribute to the public good is that the blenny may specialise on non‐punishing ‘free‐riders’. Here, we investigated how far a spatial structure within the shoal and limited space use by the blenny may contribute to punishment being self‐serving. Field observation reveals that anthias indeed live in spatially structured groups and that blennies have preferred areas for attacks. Thus, some anthias individuals are consistently more exposed to blennies than others and hence may gain disproportional benefits from their punishment. In conclusion, spatial structure may contribute to the maintenance of punishment in blenny–anthias interactions even when groups are large.