Higher frequency of social learning in China than in the West shows cultural variation in the dynamics of cultural evolution

Cultural evolutionary models have identified a range of conditions under which social learning (copying others) is predicted to be adaptive relative to asocial learning (learning on one''s own), particularly in humans where socially learned information can accumulate over successive generations. However, cultural evolution and behavioural economics experiments have consistently shown apparently maladaptive under-utilization of social information in Western populations. Here we provide experimental evidence of cultural variation in people''s use of social learning, potentially explaining this mismatch. People in mainland China showed significantly more social learning than British people in an artefact-design task designed to assess the adaptiveness of social information use. People in Hong Kong, and Chinese immigrants in the UK, resembled British people in their social information use, suggesting a recent shift in these groups from social to asocial learning due to exposure to Western culture. Finally, Chinese mainland participants responded less than other participants to increased environmental change within the task. Our results suggest that learning strategies in humans are culturally variable and not genetically fixed, necessitating the study of the ‘social learning of social learning strategies'' whereby the dynamics of cultural evolution are responsive to social processes, such as migration, education and globalization.     

Inferring population histories using cultural data

The question as to whether cultures evolve in a manner analogous to that of genetic evolution can be addressed by attempting to reconstruct population histories using cultural data. As others have argued, this can only succeed if cultures are isolated enough to maintain and pass on a central core of traditions that can be modified over time. In this study we used a set of cultural data (canoe design traits from Polynesia) to look for the kinds of patterns and relationships normally found in population genetic studies. After developing new techniques to accommodate the peculiarities of cultural data, we were able to infer an ancestral region (Fiji) and a sequence of cultural origins for these Polynesian societies. In addition, we found evidence of cultural exchange, migration and a serial founder effect. Results were stronger when analyses were based on functional traits (presumably subject to natural selection and convergence) rather than symbolic or stylistic traits (probably subject to cultural selection for rapid divergence). These patterns strongly suggest that cultural evolution, while clearly affected by cultural exchange, is also subject to some of the same processes and constraints as genetic evolution.     

Review. The multiple roles of cultural transmission experiments in understanding human cultural evolution

In this paper, we explore how experimental studies of cultural transmission in adult humans can address general questions regarding the 'who, what, when and how' of human cultural transmission, and consequently inform a theory of human cultural evolution. Three methods are discussed. The transmission chain method, in which information is passed along linear chains of participants, has been used to identify content biases in cultural transmission. These concern the kind of information that is transmitted. Several such candidate content biases have now emerged from the experimental literature. The replacement method, in which participants in groups are gradually replaced or moved across groups, has been used to study phenomena such as cumulative cultural evolution, cultural group selection and cultural innovation. The closed-group method, in which participants learn in groups with no replacement, has been used to explore issues such as who people choose to learn from and when they learn culturally as opposed to individually. A number of the studies reviewed here have received relatively little attention within their own disciplines, but we suggest that these, and future experimental studies of cultural transmission that build on them, can play an important role in a broader science of cultural evolution.     

A multilevel analytical framework for studying cultural evolution in prehistoric hunter–gatherer societies

Over the past decade, a major debate has taken place on the underpinnings of cultural changes in human societies. A growing array of evidence in behavioural and evolutionary biology has revealed that social connectivity among populations and within them affects, and is affected by, culture. Yet the interplay between prehistoric hunter–gatherer social structure and cultural transmission has typically been overlooked. Interestingly, the archaeological record contains large data sets, allowing us to track cultural changes over thousands of years: they thus offer a unique opportunity to shed light on long‐term cultural transmission processes. In this review, we demonstrate how well‐developed methods for social structure analysis can increase our understanding of the selective pressures underlying cumulative culture. We propose a multilevel analytical framework that considers finer aspects of the complex social structure in which regional groups of prehistoric hunter–gatherers were embedded. We put forward predictions of cultural transmission based on local‐ and global‐level network metrics of small‐scale societies and their potential effects on cumulative culture. By bridging the gaps between network science, palaeodemography and cultural evolution, we draw attention to the use of the archaeological record to depict patterns of social interactions and transmission variability. We argue that this new framework will contribute to improving our understanding of social interaction patterns, as well as the contexts in which cultural changes occur. Ultimately, this may provide insights into the evolution of human behaviour.     

Identifying innovation in laboratory studies of cultural evolution: rates of retention and measures of adaptation

In recent years, laboratory studies of cultural evolution have become increasingly prevalent as a means of identifying and understanding the effects of cultural transmission on the form and functionality of transmitted material. The datasets generated by these studies may provide insights into the conditions encouraging, or inhibiting, high rates of innovation, as well as the effect that this has on measures of adaptive cultural change. Here we review recent experimental studies of cultural evolution with a view to elucidating the role of innovation in generating observed trends. We first consider how tasks are presented to participants, and how the corresponding conceptualization of task success is likely to influence the degree of intent underlying any deviations from perfect reproduction. We then consider the measures of interest used by the researchers to track the changes that occur as a result of transmission, and how these are likely to be affected by differing rates of retention. We conclude that considering studies of cultural evolution from the perspective of innovation provides us with valuable insights that help to clarify important differences in research designs, which have implications for the likely effects of variation in retention rates on measures of cultural adaptation.     

Social foraging: individual learning and cultural transmission of innovations

We present two stochastic models of individual and social learningthat count the number of individuals exhibiting a learned, resource-producingtrait in a group of social foragers. The novelty of our modelingresults from incorporating the empirically based assumptionthat rates of both individual and social learning should dependon the frequency of the learned trait within the group. Whenresources occur as clumps shared by group members, a naive individual'sacquisition of the skill required for clump discovery/productionshould involve opposing processes of frequency dependence. Theopportunity to learn via cultural transmission should increasewith the trait's frequency, but the opportunity for learningindividually should decrease as the trait's frequency increases.The results of the model suggest that the evolution of the capacityfor cultural transmission may be promoted in environments wherescrounging at resource clumps inhibits rates of individual learning.     

How copying affects the amount, evenness and persistence of cultural knowledge: insights from the social learning strategies tournament

Darwinian processes should favour those individuals that deploy the most effective strategies for acquiring information about their environment. We organized a computer-based tournament to investigate which learning strategies would perform well in a changing environment. The most successful strategies relied almost exclusively on social learning (here, learning a behaviour performed by another individual) rather than asocial learning, even when environments were changing rapidly; moreover, successful strategies focused learning effort on periods of environmental change. Here, we use data from tournament simulations to examine how these strategies might affect cultural evolution, as reflected in the amount of culture (i.e. number of cultural traits) in the population, the distribution of cultural traits across individuals, and their persistence through time. We found that high levels of social learning are associated with a larger amount of more persistent knowledge, but a smaller amount of less persistent expressed behaviour, as well as more uneven distributions of behaviour, as individuals concentrated on exploiting a smaller subset of behaviour patterns. Increased rates of environmental change generated increases in the amount and evenness of behaviour. These observations suggest that copying confers on cultural populations an adaptive plasticity, allowing them to respond to changing environments rapidly by drawing on a wider knowledge base.     

Sociality influences cultural complexity

Archaeological and ethnohistorical evidence suggests a link between a population''s size and structure, and the diversity or sophistication of its toolkits or technologies. Addressing these patterns, several evolutionary models predict that both the size and social interconnectedness of populations can contribute to the complexity of its cultural repertoire. Some models also predict that a sudden loss of sociality or of population will result in subsequent losses of useful skills/technologies. Here, we test these predictions with two experiments that permit learners to access either one or five models (teachers). Experiment 1 demonstrates that naive participants who could observe five models, integrate this information and generate increasingly effective skills (using an image editing tool) over 10 laboratory generations, whereas those with access to only one model show no improvement. Experiment 2, which began with a generation of trained experts, shows how learners with access to only one model lose skills (in knot-tying) more rapidly than those with access to five models. In the final generation of both experiments, all participants with access to five models demonstrate superior skills to those with access to only one model. These results support theoretical predictions linking sociality to cumulative cultural evolution.     

Introduction. Cultural transmission and the evolution of human behaviour

The articles in this theme issue seek to understand the evolutionary bases of social learning and the consequences of cultural transmission for the evolution of human behaviour. In this introductory article, we provide a summary of these articles (seven articles on the experimental exploration of cultural transmission and three articles on the role of gene-culture coevolution in shaping human behaviour) and a personal view of some promising lines of development suggested by the work summarized here.     

Macro-evolutionary studies of cultural diversity: a review of empirical studies of cultural transmission and cultural adaptation

A growing body of theoretical and empirical research has examined cultural transmission and adaptive cultural behaviour at the individual, within-group level. However, relatively few studies have tried to examine proximate transmission or test ultimate adaptive hypotheses about behavioural or cultural diversity at a between-societies macro-level. In both the history of anthropology and in present-day work, a common approach to examining adaptive behaviour at the macro-level has been through correlating various cultural traits with features of ecology. We discuss some difficulties with simple ecological associations, and then review cultural phylogenetic studies that have attempted to go beyond correlations to understand the underlying cultural evolutionary processes. We conclude with an example of a phylogenetically controlled approach to understanding proximate transmission pathways in Austronesian cultural diversity.     

Questioning the cultural evolution of altruism

The evolutionary foundations of helping among nonkin in humans have been the object of intense debates in the past decades. One thesis has had a prominent influence in this debate: the suggestion that genuine altruism, strictly defined as a form of help that comes at a net fitness cost for the benefactor, might have evolved owing to cultural transmission. The gene–culture coevolution literature is wont to claim that cultural evolution changes the selective pressures that normally act to limit the emergence of altruistic behaviours. This paper aims to recall, however, that cultural transmission yields altruism only to the extent that it relies on maladaptive mechanisms, such as conformist imitation and (in some cases) payoff‐biased transmission. This point is sometimes obscured in the literature by a confusion between genuine altruism, maladaptive by definition, and mutualistic forms of cooperation, that benefit all parties in the long run. Theories of cultural altruism do not lift the selective pressures weighing on strictly altruistic actions; they merely shift the burden of maladaptation from social cognition to cultural transmission.     

Cumulative cultural dynamics and the coevolution of cultural innovation and transmission: an ESS model for panmictic and structured populations

When individuals in a population can acquire traits through learning, each individual may express a certain number of distinct cultural traits. These traits may have been either invented by the individual himself or acquired from others in the population. Here, we develop a game theoretic model for the accumulation of cultural traits through individual and social learning. We explore how the rates of innovation, decay, and transmission of cultural traits affect the evolutionary stable (ES) levels of individual and social learning and the number of cultural traits expressed by an individual when cultural dynamics are at a steady‐state. We explore the evolution of these phenotypes in both panmictic and structured population settings. Our results suggest that in panmictic populations, the ES level of learning and number of traits tend to be independent of the social transmission rate of cultural traits and is mainly affected by the innovation and decay rates. By contrast, in structured populations, where interactions occur between relatives, the ES level of learning and the number of traits per individual can be increased (relative to the panmictic case) and may then markedly depend on the transmission rate of cultural traits. This suggests that kin selection may be one additional solution to Rogers's paradox of nonadaptive culture.     

Review. Studying cumulative cultural evolution in the laboratory

Cumulative cultural evolution is the term given to a particular kind of social learning, which allows for the accumulation of modifications over time, involving a ratchet-like effect where successful modifications are maintained until they can be improved upon. There has been great interest in the topic of cumulative cultural evolution from researchers from a wide variety of disciplines, but until recently there were no experimental studies of this phenomenon. Here, we describe our motivations for developing experimental methods for studying cumulative cultural evolution and review the results we have obtained using these techniques. The results that we describe have provided insights into understanding the outcomes of cultural processes at the population level. Our experiments show that cumulative cultural evolution can result in adaptive complexity in behaviour and can also produce convergence in behaviour. These findings lend support to ideas that some behaviours commonly attributed to natural selection and innate tendencies could in fact be shaped by cultural processes.     

Conformist learning in nine-spined sticklebacks' foraging decisions

Theoretical analyses have reported that in most circumstances where natural selection favours reliance on social learning, conformity (positive frequency-dependent social learning) is also favoured. These findings suggest that much animal social learning should involve a copy-the-majority strategy, yet there is currently surprisingly little evidence for conformist learning among animals. Here, we investigate this possibility in the nine-spined stickleback (Pungitius pungitius) by manipulating the number of demonstrator fish at two feeders, one rich and one poor, during a demonstration phase and evaluating how this affects the likelihood that the focal fish copy the demonstrators'' apparent choices. As predicted, we observed a significantly increased level of copying with increasing numbers of demonstrators at the richer of the two feeders, with copying increasing disproportionately, rather than linearly, with the proportion of demonstrators at the rich foraging patch. Control conditions with non-feeding demonstrators showed that this was not simply the result of a preference for shoaling with larger groups, implying that nine-spined sticklebacks copy in a conformist manner.     

Spatial processes and evolutionary models: a critical review

Evolution is a fundamentally population level process in which variation, drift and selection produce both temporal and spatial patterns of change. Statistical model fitting is now commonly used to estimate which kind of evolutionary process best explains patterns of change through time using models like Brownian motion, stabilizing selection (Ornstein–Uhlenbeck) and directional selection on traits measured from stratigraphic sequences or on phylogenetic trees. But these models assume that the traits possessed by a species are homogeneous. Spatial processes such as dispersal, gene flow and geographical range changes can produce patterns of trait evolution that do not fit the expectations of standard models, even when evolution at the local‐population level is governed by drift or a typical OU model of selection. The basic properties of population level processes (variation, drift, selection and population size) are reviewed and the relationship between their spatial and temporal dynamics is discussed. Typical evolutionary models used in palaeontology incorporate the temporal component of these dynamics, but not the spatial. Range expansions and contractions introduce rate variability into drift processes, range expansion under a drift model can drive directional change in trait evolution, and spatial selection gradients can create spatial variation in traits that can produce long‐term directional trends and punctuation events depending on the balance between selection strength, gene flow, extirpation probability and model of speciation. Using computational modelling that spatial processes can create evolutionary outcomes that depart from basic population‐level notions from these standard macroevolutionary models.     

The value of teaching increases with tool complexity in cumulative cultural evolution

Human cumulative cultural evolution (CCE) is recognized as a powerful ecological and evolutionary force, but its origins are poorly understood. The long-standing view that CCE requires specialized social learning processes such as teaching has recently come under question, and cannot explain why such processes evolved in the first place. An alternative, but largely untested, hypothesis is that these processes gradually coevolved with an increasing reliance on complex tools. To address this, we used large-scale transmission chain experiments (624 participants), to examine the role of different learning processes in generating cumulative improvements in two tool types of differing complexity. Both tool types increased in efficacy across experimental generations, but teaching only provided an advantage for the more complex tools. Moreover, while the simple tools tended to converge on a common design, the more complex tools maintained a diversity of designs. These findings indicate that the emergence of cumulative culture is not strictly dependent on, but may generate selection for, teaching. As reliance on increasingly complex tools grew, so too would selection for teaching, facilitating the increasingly open-ended evolution of cultural artefacts.     

Social Learning and the Maintenance of Cultural Variation: An Evolutionary Model and Data from East Africa

Human societies maintain between-group variation despite mixing of people and ideas. In order for variation to remain, migrants or their children must preferentially adopt local norms, customs, and beliefs. Yet the details of how cultural variation is maintained, despite mixing, remain unknown. This article addresses this problem by using a simple model of the evolution of cultural learning to interpret the results of a study of cultural variation in a small region of East Africa. I argue that the manner in which migrants of two diverse regions adapt to local beliefs and behavior depends on the costs and accuracy of learning in each domain. Observational studies are never definitive tests of any hypothesis, but these results suggest that conclusions about the significance of cultural learning for understanding individual attitudes and behavior depend strongly upon the domain of investigation.     

Natural pedagogy as evolutionary adaptation

We propose that the cognitive mechanisms that enable the transmission of cultural knowledge by communication between individuals constitute a system of 'natural pedagogy' in humans, and represent an evolutionary adaptation along the hominin lineage. We discuss three kinds of arguments that support this hypothesis. First, natural pedagogy is likely to be human-specific: while social learning and communication are both widespread in non-human animals, we know of no example of social learning by communication in any other species apart from humans. Second, natural pedagogy is universal: despite the huge variability in child-rearing practices, all human cultures rely on communication to transmit to novices a variety of different types of cultural knowledge, including information about artefact kinds, conventional behaviours, arbitrary referential symbols, cognitively opaque skills and know-how embedded in means-end actions. Third, the data available on early hominin technological culture are more compatible with the assumption that natural pedagogy was an independently selected adaptive cognitive system than considering it as a by-product of some other human-specific adaptation, such as language. By providing a qualitatively new type of social learning mechanism, natural pedagogy is not only the product but also one of the sources of the rich cultural heritage of our species.     

Not-so-social learning strategies

Social learning strategies (SLSs) are rules specifying the conditions in which it would be adaptive for animals to copy the behaviour of others rather than to persist with a previously established behaviour or to acquire a new behaviour through asocial learning. In behavioural ecology, cultural evolutionary theory and economics, SLSs are studied using a ‘phenotypic gambit’—from a purely functional perspective, without reference to their underlying psychological mechanisms. However, SLSs are described in these fields as if they were implemented by complex, domain-specific, genetically inherited mechanisms of decision-making. In this article, we suggest that it is time to begin investigating the psychology of SLSs, and we initiate this process by examining recent experimental work relating to three groups of strategies: copy when alternative unsuccessful, copy when model successful and copy the majority. In each case, we argue that the reported behaviour could have been mediated by domain-general and taxonomically general psychological mechanisms; specifically, by mechanisms, identified through conditioning experiments, that make associative learning selective. We also suggest experimental manipulations that could be used in future research to resolve more fully the question whether, in non-human animals, SLSs are mediated by domain-general or domain-specific psychological mechanisms.