Social learning (learning through observation or interaction with other individuals) is widespread in nature and is central to the remarkable success of humanity, yet it remains unclear why it pays to copy, and how best to do so. To address these questions we organised a computer tournament in which entrants submitted strategies specifying how to use social learning and its asocial alternative (e.g. trial-and-error) to acquire adaptive behavior in a complex environment. Most current theory predicts the emergence of mixed strategies that rely on some combination of the two types of learning. In the tournament, however, strategies that relied heavily on social learning were found to be remarkably successful, even when asocial information was no more costly than social information. Social learning proved advantageous because individuals frequently demonstrated the highest-payoff behavior in their repertoire, inadvertently filtering information for copiers. The winning strategy (discountmachine) relied exclusively on social learning, and weighted information according to the time since acquisition.Human culture is widely thought to underlie the extraordinary demographic success of our species, manifest in virtually every terrestrial habitat (1-2). Cultural processes facilitate the spread of adaptive knowledge, accumulated over generations, allowing individuals to acquire vital life skills. One of the foundations of culture is social learning -learning influenced by observation or interaction with other individuals (3) -which occurs widely, in various forms, * To whom correspondence should be addressed. ler4@st-andrews.ac.uk.One sentence summary: A computer tournament helps to explain why social learning is common in nature and why human beings happen to be so good at it. NIH Public Access Author ManuscriptScience. Author manuscript; available in PMC 2010 November 22. Published in final edited form as:Science. 2010 April 9; 328(5975): 208-213. doi:10.1126/science.1184719. NIH-PA Author ManuscriptNIH-PA Author Manuscript NIH-PA Author Manuscript across the animal kingdom (4). Yet it remains something of a mystery why it pays individuals to copy others, and how best to do this.At first sight, social learning appears advantageous because it allows individuals to avoid the costs, in terms of effort and risk, of trial-and-error learning. However, social learning can also cost time and effort, and theoretical work reveals that it can be error prone, leading individuals to acquire inappropriate or outdated information in nonuniform and changing environments (5-11). Current theory suggests that to avoid these errors individuals should be selective in when and how they use social learning, so as to balance its advantages against the risks inherent in its indiscriminate use (9). Accordingly, natural selection is expected to have favoured social learning strategies, psychological mechanisms that specify when individuals copy, and from whom they learn (12-13).These issues lie at the interface of multiple academic fields,...

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Teaching, alongside imitation, is widely thought to underlie the success of humanity by allowing high-fidelity transmission of information, skills, and technology between individuals, facilitating both cumulative knowledge gain and normative culture. Yet, it remains a mystery why teaching should be widespread in human societies but extremely rare in other animals. We explore the evolution of teaching using simple genetic models in which a single tutor transmits adaptive information to a related pupil at a cost. Teaching is expected to evolve where its costs are outweighed by the inclusive fitness benefits that result from the tutor's relatives being more likely to acquire the valuable information. We find that teaching is not favored where the pupil can easily acquire the information on its own, or through copying others, or for difficult to learn traits, where teachers typically do not possess the information to pass on to relatives. This leads to a narrow range of traits for which teaching would be efficacious, which helps to explain the rarity of teaching in nature, its unusual distribution, and its highly specific nature. Further models that allow for cumulative cultural knowledge gain suggest that teaching evolved in humans because cumulative culture renders otherwise difficult-to-acquire valuable information available to teach.

We explore the evolution of reliance on social and asocial learning using a spatially explicit stochastic model. Our analysis considers the relative merits of four evolved strategies, two pure strategies (asocial and social learning) and two conditional strategies (the "critical social learner," which learns asocially only when copying fails, and the "conditional social learner," which copies only when asocial learning fails). We find that spatial structure generates outcomes that do not always conform to the finding of earlier theoretical analyses that social learning does not enhance average individual fitness at equilibrium (Rogers' paradox). Although we describe circumstances under which the strategy of pure social learning increases the average fitness of individuals, we find that spatial structure introduces a new paradox, which is that social learning can spread even when it decreases the average fitness of individuals below that of asocial learners. We also show that the critical social learner and conditional social learner both provide solutions to the aforementioned paradoxes, although we find some conditions in which pure (random) social learning out-competes both conditional strategies. Finally, we consider the relative merits of critical and conditional social learning under various conditions. K E Y W O R D S : Culture, evolution, learning, social learning strategy, spatial model.

Cultural niche construction is a uniquely potent source of selection on human populations, and a major cause of recent human evolution. Previous theoretical analyses have not, however, explored the local effects of cultural niche construction. Here, we use spatially explicit coevolutionary models to investigate how cultural processes could drive selection on human genes by modifying local resources. We show that cultural learning, expressed in local niche construction, can trigger a process with dynamics that resemble runaway sexual selection. Under a broad range of conditions, cultural niche-constructing practices generate selection for gene-based traits and hitchhike to fixation through the build up of statistical associations between practice and trait. This process can occur even when the cultural practice is costly, or is subject to counteracting transmission biases, or the genetic trait is selected against. Under some conditions a secondary hitchhiking occurs, through which genetic variants that enhance the capability for cultural learning are also favoured by similar dynamics. We suggest that runaway cultural niche construction could have played an important role in human evolution, helping to explain why humans are simultaneously the species with the largest relative brain size, the most potent capacity for niche construction and the greatest reliance on culture.

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Niche construction is a process through which organisms alter their environments and, in doing so, influence or change the selective pressures to which they are subject. 'Cultural niche construction' refers specifically to the effect of cultural traits on the selective environments of other biological or cultural traits and may be especially important in human evolution. In addition, the relationship between population size and cultural accumulation has been the subject of extensive debate, in part because anthropological studies have demonstrated a significant association between population size and toolkit complexity in only a subset of studied cultures. Here, we review the role of cultural innovation in constructing human evolutionary niches and introduce a new model to describe the accumulation of human cultural traits that incorporates the effects of cultural niche construction. We consider the results of this model in light of available data on human toolkit sizes across populations to help elucidate the important differences between food-gathering societies and food-producing societies, in which niche construction may be a more potent force. These results support the idea that a population's relationship with its environment, represented here by cultural niche construction, should be considered alongside population size in studies of cultural complexity.This article is part of the themed issue 'Process and pattern in innovations from cells to societies'.

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.