Convergent transformation and selection in cultural evolution

In biology, natural selection is the main explanation of adaptations and it is an attractive idea to think that an analogous force could have the same role in cultural evolution. In support of this idea, all the main ingredients for natural selection have been documented in the cultural domain. However, the changes that occur during cultural transmission typically result in convergent transformation, non-random cultural modifications, casting some doubts on the importance of natural selection in the cultural domain. To progress on this issue more empirical research is needed. Here, using nearly half a million experimental trials performed by a group of baboons (Papio papio), we simulate cultural evolution under various conditions of natural selection and do an additional experiment to tease apart the role of convergent transformation and selection. Our results confirm that transformation strongly constrain the variation available to selection and therefore strongly limit its impact on cultural evolution. Surprisingly, in our study, transformation also enhances the effect of selection by stabilising cultural variation. We conclude that, in culture, selection can change the evolutionary trajectory substantially in some cases, but can only act on the variation provided by (typically biased) transformation.     

Review. Theoretical and empirical evidence for the impact of inductive biases on cultural evolution

The question of how much the outcomes of cultural evolution are shaped by the cognitive capacities of human learners has been explored in several disciplines, including psychology, anthropology and linguistics. We address this question through a detailed investigation of transmission chains, in which each person passes information to another along a chain. We review mathematical and empirical evidence that shows that under general conditions, and across experimental paradigms, the information passed along transmission chains will be affected by the inductive biases of the people involved-the constraints on learning and memory, which influence conclusions from limited data. The mathematical analysis considers the case where each person is a rational Bayesian agent. The empirical work consists of behavioural experiments in which human participants are shown to operate in the manner predicted by the Bayesian framework. Specifically, in situations in which each person's response is used to determine the data seen by the next person, people converge on concepts consistent with their inductive biases irrespective of the information seen by the first member of the chain. We then relate the Bayesian analysis of transmission chains to models of biological evolution, clarifying how chains of individuals correspond to population-level models and how selective forces can be incorporated into our models. Taken together, these results indicate how laboratory studies of transmission chains can provide information about the dynamics of cultural evolution and illustrate that inductive biases can have a significant impact on these dynamics.     

Five fundamental constraints on theories of the origins of music

The diverse forms and functions of human music place obstacles in the way of an evolutionary reconstruction of its origins. In the absence of any obvious homologues of human music among our closest primate relatives, theorizing about its origins, in order to make progress, needs constraints from the nature of music, the capacities it engages, and the contexts in which it occurs. Here we propose and examine five fundamental constraints that bear on theories of how music and some of its features may have originated. First, cultural transmission, bringing the formal powers of cultural as contrasted with Darwinian evolution to bear on its contents. Second, generativity, i.e. the fact that music generates infinite pattern diversity by finite means. Third, vocal production learning, without which there can be no human singing. Fourth, entrainment with perfect synchrony, without which there is neither rhythmic ensemble music nor rhythmic dancing to music. And fifth, the universal propensity of humans to gather occasionally to sing and dance together in a group, which suggests a motivational basis endemic to our biology. We end by considering the evolutionary context within which these constraints had to be met in the genesis of human musicality.     

A model for the evolutionary diversification of religions

We address the problem of cultural diversification by studying selection on cultural ideas that colonize human hosts and using diversification of religions as a conceptual example. In analogy to studying the evolution of pathogens or symbionts colonizing animal hosts, we use models for host-pathogen dynamics known from theoretical epidemiology. In these models, religious content colonizes individual humans. Rates of transmission of ideas between humans, i.e., transmission of cultural content, and rates of loss of ideas (loss of belief) are determined by the phenotype of the cultural content, and by interactions between hosts carrying different ideas. In particular, based on the notion that cultural non-conformism can be negative frequency-dependent (for example, religion can lead to oppression of lower classes and emergence of non-conformism and dissent once a religious belief has reached dominance), we assume that the rate of loss of belief increases as the number of humans colonized by a particular religious phenotype increases. This generates frequency-dependent selection on cultural content, and we use evolutionary theory to show that this frequency dependence can lead to the emergence of coexisting clusters of different cultural types. The different clusters correspond to different cultural traditions, and hence our model describes the emergence of distinct descendant cultures from a single ancestral culture in the absence of any geographical isolation.     

Big brains, small worlds: material culture and the evolution of the mind

New developments in neuroimaging have demonstrated that the basic capacities underpinning human social skills are shared by our closest extant primate relatives. The challenge for archaeologists is to explain how complex human societies evolved from this shared pattern of face-to-face social interaction. We argue that a key process was the gradual incorporation of material culture into social networks over the course of hominin evolution. Here we use three long-term processes in hominin evolution-encephalization, the global human diaspora and sedentism/agriculture-to illustrate how the cultural transmission of material culture allowed the 'scaling up' of face-to-face social interactions to the global societies known today. We conclude that future research by neuroimagers and archaeologists will need to investigate the cognitive mechanisms behind human engagement with material culture as well as other persons.     

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.     

Cognitive capacities for cooking in chimpanzees

The transition to a cooked diet represents an important shift in human ecology and evolution. Cooking requires a set of sophisticated cognitive abilities, including causal reasoning, self-control and anticipatory planning. Do humans uniquely possess the cognitive capacities needed to cook food? We address whether one of humans'' closest relatives, chimpanzees (Pan troglodytes), possess the domain-general cognitive skills needed to cook. Across nine studies, we show that chimpanzees: (i) prefer cooked foods; (ii) comprehend the transformation of raw food that occurs when cooking, and generalize this causal understanding to new contexts; (iii) will pay temporal costs to acquire cooked foods; (iv) are willing to actively give up possession of raw foods in order to transform them; and (v) can transport raw food as well as save their raw food in anticipation of future opportunities to cook. Together, our results indicate that several of the fundamental psychological abilities necessary to engage in cooking may have been shared with the last common ancestor of apes and humans, predating the control of fire.     

The evolution of primate general and cultural intelligence

There are consistent individual differences in human intelligence, attributable to a single 'general intelligence' factor, g. The evolutionary basis of g and its links to social learning and culture remain controversial. Conflicting hypotheses regard primate cognition as divided into specialized, independently evolving modules versus a single general process. To assess how processes underlying culture relate to one another and other cognitive capacities, we compiled ecologically relevant cognitive measures from multiple domains, namely reported incidences of behavioural innovation, social learning, tool use, extractive foraging and tactical deception, in 62 primate species. All exhibited strong positive associations in principal component and factor analyses, after statistically controlling for multiple potential confounds. This highly correlated composite of cognitive traits suggests social, technical and ecological abilities have coevolved in primates, indicative of an across-species general intelligence that includes elements of cultural intelligence. Our composite species-level measure of general intelligence, 'primate g(S)', covaried with both brain volume and captive learning performance measures. Our findings question the independence of cognitive traits and do not support 'massive modularity' in primate cognition, nor an exclusively social model of primate intelligence. High general intelligence has independently evolved at least four times, with convergent evolution in capuchins, baboons, macaques and great apes.     

Societal background influences social learning in cooperative decision making

Humans owe their ecological success to their great capacities for social learning and cooperation: learning from others helps individuals adjust to their environment and can promote cooperation in groups. Classic and recent studies indicate that the cultural organization of societies shapes the influence of social information on decision making and suggest that collectivist values (prioritizing the group relative to the individual) increase tendencies to conform to the majority. However, it is unknown whether and how societal background impacts social learning in cooperative interactions. Here we show that social learning in cooperative decision making systematically varies across two societies. We experimentally compare people's basic propensities for social learning in samples from a collectivist (China) and an individualist society (United Kingdom; total n?=?540) in a social dilemma and a coordination game. We demonstrate that Chinese participants base their cooperation decisions on information about their peers much more frequently than their British counterparts. Moreover, our results reveal remarkable societal differences in the type of peer information people consider. In contrast to the consensus view, Chinese participants tend to be substantially less majority-oriented than the British. While Chinese participants are inclined to adopt peer behavior that leads to higher payoffs, British participants tend to cooperate only if sufficiently many peers do so too. These results indicate that the basic processes underlying social transmission are not universal; rather, they vary with cultural conditions. As success-based learning is associated with selfish behavior and majority-based learning can help foster cooperation, our study suggests that in different societies social learning can play diverging roles in the emergence and maintenance of cooperation.     

Innovation in the collective brain

Innovation is often assumed to be the work of a talented few, whose products are passed on to the masses. Here, we argue that innovations are instead an emergent property of our species'' cultural learning abilities, applied within our societies and social networks. Our societies and social networks act as collective brains. We outline how many human brains, which evolved primarily for the acquisition of culture, together beget a collective brain. Within these collective brains, the three main sources of innovation are serendipity, recombination and incremental improvement. We argue that rates of innovation are heavily influenced by (i) sociality, (ii) transmission fidelity, and (iii) cultural variance. We discuss some of the forces that affect these factors. These factors can also shape each other. For example, we provide preliminary evidence that transmission efficiency is affected by sociality—languages with more speakers are more efficient. We argue that collective brains can make each of their constituent cultural brains more innovative. This perspective sheds light on traits, such as IQ, that have been implicated in innovation. A collective brain perspective can help us understand otherwise puzzling findings in the IQ literature, including group differences, heritability differences and the dramatic increase in IQ test scores over time.     

Why does human culture increase exponentially?

Historical records show that culture can increase exponentially in time, e.g., in number of poems, musical works, scientific discoveries. We model how human capacities for creativity and cultural transmission may make such an increase possible, suggesting that: (1) creativity played a major role at the origin of human culture and for its accumulation throughout history, because cultural transmission cannot, on its own, generate exponentially increasing amounts of culture; (2) exponential increase in amount of culture can only occur if creativity is positively influenced by culture. The evolution of cultural transmission is often considered the main genetic bottleneck for the origin of culture, because natural selection cannot favor cultural transmission without any culture to transmit. Our models suggest that an increase in individual creativity may have been the first step toward human culture, because in a population of creative individuals there may be enough non-genetic information to favor the evolution of cultural transmission.     

Culture evolves

Culture pervades human lives and has allowed our species to create niches all around the world and its oceans, in ways quite unlike any other primate. Indeed, our cultural nature appears so distinctive that it is often thought to separate humanity from the rest of nature and the Darwinian forces that shape it. A contrary view arises through the recent discoveries of a diverse range of disciplines, here brought together to illustrate the scope of a burgeoning field of cultural evolution and to facilitate cross-disciplinary fertilization. Each approach emphasizes important linkages between culture and evolutionary biology rather than quarantining one from the other. Recent studies reveal that processes important in cultural transmission are more widespread and significant across the animal kingdom than earlier recognized, with important implications for evolutionary theory. Recent archaeological discoveries have pushed back the origins of human culture to much more ancient times than traditionally thought. These developments suggest previously unidentified continuities between animal and human culture. A third new array of discoveries concerns the later diversification of human cultures, where the operations of Darwinian-like processes are identified, in part, through scientific methods borrowed from biology. Finally, surprising discoveries have been made about the imprint of cultural evolution in the predispositions of human minds for cultural transmission.     

Evolution of social learning does not explain the origin of human cumulative culture

Because culture requires transmission of information between individuals, thinking about the origin of culture has mainly focused on the genetic evolution of abilities for social learning. Current theory considers how social learning affects the adaptiveness of a single cultural trait, yet human culture consists of the accumulation of very many traits. Here we introduce a new modeling strategy that tracks the adaptive value of many cultural traits, showing that genetic evolution favors only limited social learning owing to the accumulation of maladaptive as well as adaptive culture. We further show that culture can be adaptive, and refined social learning can evolve, if individuals can identify and discard maladaptive culture. This suggests that the evolution of such "adaptive filtering" mechanisms may have been crucial for the birth of human culture.     

Adaptive Social Learning Strategies in Temporally and Spatially Varying Environments

Long before the origins of agriculture human ancestors had expanded across the globe into an immense variety of environments, from Australian deserts to Siberian tundra. Survival in these environments did not principally depend on genetic adaptations, but instead on evolved learning strategies that permitted the assembly of locally adaptive behavioral repertoires. To develop hypotheses about these learning strategies, we have modeled the evolution of learning strategies to assess what conditions and constraints favor which kinds of strategies. To build on prior work, we focus on clarifying how spatial variability, temporal variability, and the number of cultural traits influence the evolution of four types of strategies: (1) individual learning, (2) unbiased social learning, (3) payoff-biased social learning, and (4) conformist transmission. Using a combination of analytic and simulation methods, we show that spatial??but not temporal??variation strongly favors the emergence of conformist transmission. This effect intensifies when migration rates are relatively high and individual learning is costly. We also show that increasing the number of cultural traits above two favors the evolution of conformist transmission, which suggests that the assumption of only two traits in many models has been conservative. We close by discussing how (1) spatial variability represents only one way of introducing the low-level, nonadaptive phenotypic trait variation that so favors conformist transmission, the other obvious way being learning errors, and (2) our findings apply to the evolution of conformist transmission in social interactions. Throughout we emphasize how our models generate empirical predictions suitable for laboratory testing.     

The evolution of cultural adaptations: Fijian food taboos protect against dangerous marine toxins

The application of evolutionary theory to understanding the origins of our species'' capacities for social learning has generated key insights into cultural evolution. By focusing on how our psychology has evolved to adaptively extract beliefs and practices by observing others, theorists have hypothesized how social learning can, over generations, give rise to culturally evolved adaptations. While much field research documents the subtle ways in which culturally transmitted beliefs and practices adapt people to their local environments, and much experimental work reveals the predicted patterns of social learning, little research connects real-world adaptive cultural traits to the patterns of transmission predicted by these theories. Addressing this gap, we show how food taboos for pregnant and lactating women in Fiji selectively target the most toxic marine species, effectively reducing a woman''s chances of fish poisoning by 30 per cent during pregnancy and 60 per cent during breastfeeding. We further analyse how these taboos are transmitted, showing support for cultural evolutionary models that combine familial transmission with selective learning from locally prestigious individuals. In addition, we explore how particular aspects of human cognitive processes increase the frequency of some non-adaptive taboos. This case demonstrates how evolutionary theory can be deployed to explain both adaptive and non-adaptive behavioural patterns.     

The Big Man Mechanism: how prestige fosters cooperation and creates prosocial leaders

Anthropological evidence from diverse societies suggests that prestige-based leadership may provide a foundation for cooperation in many contexts. Here, inspired by such ethnographic observations and building on a foundation of existing research on the evolution of prestige, we develop a set of formal models to explore when an evolved prestige psychology might drive the cultural evolution of n-person cooperation, and how such a cultural evolutionary process might create novel selection pressures for genes that make prestigious individuals more prosocial. Our results reveal (i) how prestige can foster the cultural emergence of cooperation by generating correlated behavioural phenotypes, both between leaders and followers, and among followers; (ii) why, in the wake of cultural evolution, natural selection favours genes that make prestigious leaders more prosocial, but only when groups are relatively small; and (iii), why the effectiveness of status differences in generating cooperation in large groups depends on cultural transmission (and not primarily on deference or coercion). Our theoretical framework, and the specific predictions made by these models, sketch out an interdisciplinary research programme that cross-cuts anthropology, biology, psychology and economics. Some of our predictions find support from laboratory work in behavioural economics and are consistent with several real-world patterns.     

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.     

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.     

Modeling Coevolution between Language and Memory Capacity during Language Origin

Memory is essential to many cognitive tasks including language. Apart from empirical studies of memory effects on language acquisition and use, there lack sufficient evolutionary explorations on whether a high level of memory capacity is prerequisite for language and whether language origin could influence memory capacity. In line with evolutionary theories that natural selection refined language-related cognitive abilities, we advocated a coevolution scenario between language and memory capacity, which incorporated the genetic transmission of individual memory capacity, cultural transmission of idiolects, and natural and cultural selections on individual reproduction and language teaching. To illustrate the coevolution dynamics, we adopted a multi-agent computational model simulating the emergence of lexical items and simple syntax through iterated communications. Simulations showed that: along with the origin of a communal language, an initially-low memory capacity for acquired linguistic knowledge was boosted; and such coherent increase in linguistic understandability and memory capacities reflected a language-memory coevolution; and such coevolution stopped till memory capacities became sufficient for language communications. Statistical analyses revealed that the coevolution was realized mainly by natural selection based on individual communicative success in cultural transmissions. This work elaborated the biology-culture parallelism of language evolution, demonstrated the driving force of culturally-constituted factors for natural selection of individual cognitive abilities, and suggested that the degree difference in language-related cognitive abilities between humans and nonhuman animals could result from a coevolution with language.