Social enhancement can create adaptive,arbitrary and maladaptive cultural traditions

Many animals are known to learn socially, i.e. they are able to acquire new behaviours by using information from other individuals. Researchers distinguish between a number of different social-learning mechanisms such as imitation and social enhancement. Social enhancement is a simple form of social learning that is among the most widespread in animals. However, unlike imitation, it is debated whether social enhancement can create cultural traditions. Based on a recent study on capuchin monkeys, we developed an agent-based model to test the hypotheses that (i) social enhancement can create and maintain stable traditions and (ii) social enhancement can create cultural conformity. Our results supported both hypotheses. A key factor that led to the creation of cultural conformity and traditions was the repeated interaction of individual reinforcement and social enhancement learning. This result emphasizes that the emergence of cultural conformity does not necessarily require cognitively complex mechanisms such as ‘copying the majority’ or group norms. In addition, we observed that social enhancement can create learning dynamics similar to a ‘copy when uncertain’ learning strategy. Results from additional analyses also point to situations that should favour the evolution of learning mechanisms more sophisticated than social enhancement.     

Automatic imitation in budgerigars

A fully automated procedure, involving computer-controlled stimulus presentation and computer-recorded response measurement, was used for the first time to study imitation in non-human animals. After preliminary training to peck and step on a manipulandum, budgerigars were given a discrimination task in which they were rewarded with food for pecking during observation of pecking and for stepping during observation of stepping (Compatible group), or for pecking while observing stepping and for stepping while observing pecking (Incompatible group). The Incompatible group, which had to counter-imitate for food reward, showed weaker discrimination performance than the Compatible group. This suggests that, like humans, budgerigars are subject to 'automatic imitation'; they cannot inhibit online the tendency to imitate pecking and/or stepping, even when imitation of these behaviours interferes with the performance of an ongoing task. The difference between the two groups persisted over 10 test sessions, but the Incompatible group eventually acquired the discrimination, making more counter-imitative than imitative responses in the final sessions. These results are consistent with the associative sequence learning model, which suggests that, across species, the development of imitation and the mirror system depends on sensorimotor experience and phylogenetically ancient mechanisms of associative learning.     

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.     

Investigating the Relationship between Stable Personality Characteristics and Automatic Imitation

Automatic imitation is a cornerstone of nonverbal communication that fosters rapport between interaction partners. Recent research has suggested that stable dimensions of personality are antecedents to automatic imitation, but the empirical evidence linking imitation with personality traits is restricted to a few studies with modest sample sizes. Additionally, atypical imitation has been documented in autism spectrum disorders and schizophrenia, but the mechanisms underpinning these behavioural profiles remain unclear. Using a larger sample than prior studies (N=243), the current study tested whether performance on a computer-based automatic imitation task could be predicted by personality traits associated with social behaviour (extraversion and agreeableness) and with disorders of social cognition (autistic-like and schizotypal traits). Further personality traits (narcissism and empathy) were assessed in a subsample of participants (N=57). Multiple regression analyses showed that personality measures did not predict automatic imitation. In addition, using a similar analytical approach to prior studies, no differences in imitation performance emerged when only the highest and lowest 20 participants on each trait variable were compared. These data weaken support for the view that stable personality traits are antecedents to automatic imitation and that neural mechanisms thought to support automatic imitation, such as the mirror neuron system, are dysfunctional in autism spectrum disorders or schizophrenia. In sum, the impact that personality variables have on automatic imitation is less universal than initial reports suggest.     

Social facilitation of exploratory foraging behavior in capuchin monkeys (Cebus apella)

Much of the research on animal social learning focuses on complex cognitive functions such as imitation and emulation. When compelling evidence for such processes is not forthcoming, simpler processes are often assumed but rarely directly tested for. In this study we address the phenomenon of social facilitation, whereby the presence of a feeding conspecific is hypothesized to affect the motivation and behavior of the subject, elevating the likelihood of exploration and discovery in relation to the task at hand. Using a novel foraging task, sufficiently challenging that only just over half the subjects successfully gained food from it, we compared the performance of capuchin monkeys working either alone, or in a “social” condition where an actively feeding conspecific was in an adjacent chamber. Although similar numbers of subjects in these conditions were eventually successful during the 20 trials presented, the latency to successful solution of the task was over three times faster for monkeys in the social condition. The minority of monkeys that failed to learn (9/23) were then exposed to a proficient model. Only those older than 5 years provided evidence of learning from this. Accordingly, we obtained evidence for the social facilitation the study was designed to test for, and limited supplementary evidence for social learning in the older individuals who had not learned individually. These results are discussed in relation to other recent evidence for social learning in monkeys. Am. J. Primatol. 71:419–426, 2009. © 2009 Wiley‐Liss, Inc.     

Social processes affecting the appearance of innovative behaviors in capuchin monkeys

We consider the role of social factors in the appearance of innovative problem-solving behaviors in capuchins. Processes supporting dissemination of innovative behaviors included social enhancement of interest in the objects and area around the task and (rarely) coaction. Factors inhibiting display of innovative behaviors included restricted access and requirements for social vigilance among animals of vulnerable social status and exploitation by noninnovators of others' behaviors. We urge caution in interpreting the increased appearance of novel behaviors in primates as evidence for imitation or facilitation and call for a more rigorous experimental investigation of social learning in primates.     

Untersuchung zum Einfluß des Erkundungs- und Spielverhaltens auf das Lernen bei Iltisfrettchen (Mustela putorius X M. furo)

This paper investigates the dependence of learning performance on previous exploration and playing with objects. The subjects are polecats (Mustela putorius X M. furo). Behaviour of social animals, who are offered changing tube systems in certain periods, is described and analysed. A maze-test shows that subjects with tube training are superior to subjects without tube training as regards acquisition and reversal learning. The learning performance in pattern recognition tests is the same in all tests with the different groups, however.     

Context-dependent model-based biases in cultural transmission: children's imitation is affected by model age over model knowledge state

Many animals, including humans, acquire information through social learning. Although such information can be acquired easily, its potential unreliability means it should not be used indiscriminately. Cultural ‘transmission biases’ may allow individuals to weigh their reliance on social information according to a model's characteristics. In one of the first studies to juxtapose two model-based biases, we investigated whether the age and knowledge state of a model affected the fidelity of children's copying. Eighty-five 5-year-old children watched a video demonstration of either an adult or child, who had professed either knowledge or ignorance regarding a tool-use task, extracting a reward from that task using both causally relevant and irrelevant actions. Relevant actions were imitated faithfully by children regardless of the model's characteristics, but children who observed an adult reproduced more irrelevant actions than those who observed a child. The professed knowledge state of the model showed a weaker effect on imitation of irrelevant actions. Overall, children favored the use of a ‘copy adults’ bias over a ‘copy task-knowledgeable individual’ bias, even though the latter could potentially have provided more reliable information. The use of such social learning strategies has significant implications for understanding the phenomenon of imitation of irrelevant actions (overimitation), instances of maladaptive information cascades, and cumulative culture.     

The actions of others act as a pseudo-reward to drive imitation in the context of social reinforcement learning

While there is no doubt that social signals affect human reinforcement learning, there is still no consensus about how this process is computationally implemented. To address this issue, we compared three psychologically plausible hypotheses about the algorithmic implementation of imitation in reinforcement learning. The first hypothesis, decision biasing (DB), postulates that imitation consists in transiently biasing the learner’s action selection without affecting their value function. According to the second hypothesis, model-based imitation (MB), the learner infers the demonstrator’s value function through inverse reinforcement learning and uses it to bias action selection. Finally, according to the third hypothesis, value shaping (VS), the demonstrator’s actions directly affect the learner’s value function. We tested these three hypotheses in 2 experiments (N = 24 and N = 44) featuring a new variant of a social reinforcement learning task. We show through model comparison and model simulation that VS provides the best explanation of learner’s behavior. Results replicated in a third independent experiment featuring a larger cohort and a different design (N = 302). In our experiments, we also manipulated the quality of the demonstrators’ choices and found that learners were able to adapt their imitation rate, so that only skilled demonstrators were imitated. We proposed and tested an efficient meta-learning process to account for this effect, where imitation is regulated by the agreement between the learner and the demonstrator. In sum, our findings provide new insights and perspectives on the computational mechanisms underlying adaptive imitation in human reinforcement learning.

This study investigates imitation from a computational perspective; three experiments show that, in the context of reinforcement learning, imitation operates via a durable modification of the learner''s values, shedding new light on how imitation is computationally implemented and shapes learning and decision-making.     

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.     

Imitation as behaviour parsing

Non-human great apes appear to be able to acquire elaborate skills partly by imitation, raising the possibility of the transfer of skill by imitation in animals that have only rudimentary mentalizing capacities: in contrast to the frequent assumption that imitation depends on prior understanding of others' intentions. Attempts to understand the apes' behaviour have led to the development of a purely mechanistic model of imitation, the 'behaviour parsing' model, in which the statistical regularities that are inevitable in planned behaviour are used to decipher the organization of another agent's behaviour, and thence to imitate parts of it. Behaviour can thereby be understood statistically in terms of its correlations (circumstances of use, effects on the environment) without understanding of intentions or the everyday physics of cause-and-effect. Thus, imitation of complex, novel behaviour may not require mentalizing, but conversely behaviour parsing may be a necessary preliminary to attributing intention and cause.     

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.     

Success-biased social learning: Cultural and evolutionary dynamics

Success bias is a social learning strategy whereby learners tend to acquire the cultural variants of successful individuals. I develop a general model of success-biased social learning for discrete cultural traits with stochastic payoffs, and investigate its dynamics when only two variants are present. I find that success bias inherently favors rare variants, and consequently performs worse than unbiased imitation (i.e. random copying) when success payoffs are at least mildly stochastic and the optimal variant is common. Because of this weakness, success bias fails to replace unbiased imitation in an evolutionary model when selection is fairly weak or when the environment is relatively stable, and sometimes fails to invade at all. I briefly discuss the optimal strength of success bias, the complicated nature of defining success in social learning contexts, and the value of variant frequency as an important source of information to social learners. I conclude with predictions regarding the prevalence of success bias in different behavioral domains.     

Social stimuli as incentives for delayed response performance by infant pigtailed macaques (Macaca nemestrina)

Six pigtailed macaque monkeys (Macaca nemestrina), ranging in age from 4.5 to 13.5 months, were studied longitudinally on a delayed response procedure in a spatial choice apparatus. In each trial the subjects were exposed to two stimulus animals, an unfamiliar adult female and a familiar age-mate. Prior work showed that such nursery-reared infants overwhelmingly prefer a familiar peer over an unfamiliar adult female. Therefore, the peer was considered to be a positive incentive and choosing the peer was defined as the “correct” response. After the infants were given visual exposure to the stimulus animals, an opaque door was lowered to block them from the subject's view. Then a single delay period of 0, 5, 15, 60, 120, 240, or 480 sec was introduced during 20-trial sessions. The delays were increased over sessions, with about two weeks between sessions. The subjects reliably chose the familiar peer after delays of up to 60 sec, with one subject maintaining correct choices even after 8-min delays. These results revealed that the delayed response performance of young macaques with social incentives was as good as, or better than, maximum performance levels reported for macaques with food incentives. Cues such as odors, sounds, and visual-postural orientation sets could not explain the performance levels at long delays found in this study.     

Imitative behavior. A theoretical view

This article reviews the results of experimental studies on imitative behavior reported by various investigators, and then discusses the possible brain mechanisms responsible for this behavior. It was found that human infants in their first hours of life were already capable of spontaneous imitation of simple motor acts demonstrated by an adult, without previous training or reward; these observations suggest that imitative behavior is an innate process that can be considered an unconditional reflex of imitation. It was also found that satiated animals resumed eating when they saw their companions eating. In the latter case, the imitative reflex triggered the previously acquired feeding behavior. Similar mechanisms could be responsible for the phenomenon of eating more in the presence of companions than in their absence, as well as that of preferring the food chosen by companions. When followed by a reward, the imitative act can be learned--that is, transformed into an instrumental conditional response; learning by imitation of simple motor acts was observed in animals, and that of complex motor acts was observed in children who had already achieved a certain developmental stage. In animals, learning complex motor tasks was facilitated by previous observation of a companion performing this task. In this case, the presence of the observer during the session could lead to habituation of the experimental situation and production of associations between this situation and stimuli or emotions related to the reward or punishment, and might result in more efficient learning later. The imitative behavior can be inhibited by stimuli producing responses antagonistic to the act of imitation.     

How does the reliability of a model affect children's choice to learn socially or individually?

The effect of model reliability on children's choices to learn socially versus individually is pertinent to theories addressing cultural evolution and theories of selective trust. Here the effect of a reliable versus unreliable model on children's preferences to learn socially or individually was examined, as well as their subsequent imitation on a puzzle box task. Experiment One (N = 156) found children were more likely to ask to learn socially when presented with a novel task, after witnessing an unreliable rather than a reliable model. Experiment Two (N = 40) found children select a new unknown model, over the previously unreliable model, suggesting a preference to learn socially was created, although not specifically from the unreliable model. Experiment Three (N = 48) replicated children's learning preference in Experiment One with a new task, and showed children's attention is drawn towards other sources of social information (another adult model) when viewing an unreliable model, and also found a reliable model caused more fidelity of imitation. Together these results suggest that model unreliability causes greater social learning requests and attention to other, even novel, models when they are available. These findings evidence human children's strong propensity to learn socially compared with non-human animals; and suggest there is a more complicated relationship between learning preference, model reliability and selective trust than has been captured in previous research.     

Task-specific effects of ostracism on imitative fidelity in early childhood

This study examined task specific effects of third-party ostracism on imitative fidelity in early childhood (N = 96, 3–6-year-olds). Start- and end-states of action sequences were manipulated to examine the effects of priming third-party ostracism versus affiliation on children's imitation of instrumental (i.e., action sequence with a different start- and end-state) versus social convention (i.e., action sequence with an identical start- and end-state) tasks. Children's performance was coded for imitative fidelity and children's explanations for their behavior. As predicted, imitative fidelity was highest and social convention explanations were most common when primed with ostracism in the social convention task. The data are consistent with our proposal that imitation serves an affiliative function in response to the threat of ostracism, a response amplified for social conventions.     

Social learning among Congo Basin hunter-gatherers

This paper explores childhood social learning among Aka and Bofi hunter-gatherers in Central Africa. Existing literature suggests that hunter-gatherer social learning is primarily vertical (parent-to-child) and that teaching is rare. We use behavioural observations, open-ended and semi-structured interviews, and informal and anecdotal observations to examine the modes (e.g. vertical versus horizontal/oblique) and processes (e.g. teaching versus observation and imitation) of cultural transmission. Cultural and demographic contexts of social learning associated with the modes and processes of cultural transmission are described. Hunter-gatherer social learning occurred early, was relatively rapid, primarily vertical under age 5 and oblique and horizontal between the ages of 6 and 12. Pedagogy and other forms of teaching existed as early as 12 months of age, but were relatively infrequent by comparison to other processes of social learning such as observation and imitation.     

Hypotheses on mechanisms underlying observational learning in animals

Learning through observation or vicarious learning has been systematically studied in a variety of animal species for only 20 years. Demonstrating in animals a capacity to benefit from a conspecific's experience, this type of acquisition was first thought to require superior cognitive processes and thus to be restricted to primates, if possible at all in animals. The concept of imitation was commonly applied in this line of study to any social transmission. Later, experiments on vicarious learning showed that numerous species (rodents, cats, birds, primates, etc.) learn more quickly to perform an act whenever they have the opportunity of watching a conspecific performing that act. The principal characteristic of this acquisition is that it occurs during the observation period when the observer has no opportunity of either performing a response or receiving reinforcement. Four hypothesis have been put forward to explain the mechanisms underlying this type of acquisition : local enhancement, the opportunity for mediate responses, a sensory preconditioning, and the monitoring of the observer's response by some quantitative and/or qualitative aspects of the model's response. The fact that animals prove to have such a capacity is moreover a factor to be kept in mind in all studies in the eco-ethological field (particularly those concerning the predator-prey relation).