Large-scale cooperation in small-scale foraging societies.

We present evidence that people in small-scale mobile hunter-gatherer societies cooperated in large numbers to produce collective goods. Foragers engaged in large-scale communal hunts and constructed shared capital facilities; they made shared investments in improving the local environment; and they participated in warfare, formed enduring alliances, and established trading networks. Large-scale collective action often played a crucial role in subsistence. The provision of public goods involved the cooperation of many individuals, so each person made only a small contribution. This evidence suggests that large-scale cooperation occurred in the Pleistocene societies that encompass most of human evolutionary history, and therefore it is unlikely that large-scale cooperation in Holocene food producing societies results from an evolved psychology shaped only in small-group interactions. Instead, large-scale human cooperation needs to be explained as an adaptation, likely rooted in distinctive features of human biology, grammatical language, increased cognitive ability, and cumulative cultural adaptation.

Genetic solutions to cultural problems?

In theory, observed correlations between genetic information and behaviour might be useful to members of the WEIRD (western, educated, industrialized, rich, and democratic) populations. Guiding young people to choose educational opportunities that best match their abilities would benefit both the individual and society. In practice, however, such choices are far more profoundly limited by the culture people have inherited than their genes.

Collective action and the evolution of social norm internalization.

Human behavior is strongly affected by culturally transmitted norms and values. Certain norms are internalized (i.e., acting according to a norm becomes an end in itself rather than merely a tool in achieving certain goals or avoiding social sanctions). Humans' capacity to internalize norms likely evolved in our ancestors to simplify solving certain challenges-including social ones. Here we study theoretically the evolutionary origins of the capacity to internalize norms. In our models, individuals can choose to participate in collective actions as well as punish free riders. In making their decisions, individuals attempt to maximize a utility function in which normative values are initially irrelevant but play an increasingly important role if the ability to internalize norms emerges. Using agent-based simulations, we show that norm internalization evolves under a wide range of conditions so that cooperation becomes "instinctive." Norm internalization evolves much more easily and has much larger effects on behavior if groups promote peer punishment of free riders. Promoting only participation in collective actions is not effective. Typically, intermediate levels of norm internalization are most frequent but there are also cases with relatively small frequencies of "oversocialized" individuals willing to make extreme sacrifices for their groups no matter material costs, as well as "undersocialized" individuals completely immune to social norms. Evolving the ability to internalize norms was likely a crucial step on the path to large-scale human cooperation.

A theory limited in scope and evidence.

What promised to be a refreshing addition to cumulative cultural evolution, by moving the focus from cultural transmission to technological innovation, falls flat through a lack of thoroughness, explanatory power, and data. A comprehensive theory of cumulative cultural change must carefully integrate all existing evidence in a cohesive multi-level account. We argue that the manuscript fails to do so convincingly.

Current perspectives and the future of domestication studies.

It is difficult to overstate the cultural and biological impacts that the domestication of plants and animals has had on our species. Fundamental questions regarding where, when, and how many times domestication took place have been of primary interest within a wide range of academic disciplines. Within the last two decades, the advent of new archaeological and genetic techniques has revolutionized our understanding of the pattern and process of domestication and agricultural origins that led to our modern way of life. In the spring of 2011, 25 scholars with a central interest in domestication representing the fields of genetics, archaeobotany, zooarchaeology, geoarchaeology, and archaeology met at the National Evolutionary Synthesis Center to discuss recent domestication research progress and identify challenges for the future. In this introduction to the resulting Special Feature, we present the state of the art in the field by discussing what is known about the spatial and temporal patterns of domestication, and controversies surrounding the speed, intentionality, and evolutionary aspects of the domestication process. We then highlight three key challenges for future research. We conclude by arguing that although recent progress has been impressive, the next decade will yield even more substantial insights not only into how domestication took place, but also when and where it did, and where and why it did not.

Storytelling and story testing in domestication.

The domestication of plants and animals marks one of the most significant transitions in human, and indeed global, history. Traditionally, study of the domestication process was the exclusive domain of archaeologists and agricultural scientists; today it is an increasingly multidisciplinary enterprise that has come to involve the skills of evolutionary biologists and geneticists. Although the application of new information sources and methodologies has dramatically transformed our ability to study and understand domestication, it has also generated increasingly large and complex datasets, the interpretation of which is not straightforward. In particular, challenges of equifinality, evolutionary variance, and emergence of unexpected or counter-intuitive patterns all face researchers attempting to infer past processes directly from patterns in data. We argue that explicit modeling approaches, drawing upon emerging methodologies in statistics and population genetics, provide a powerful means of addressing these limitations. Modeling also offers an approach to analyzing datasets that avoids conclusions steered by implicit biases, and makes possible the formal integration of different data types. Here we outline some of the modeling approaches most relevant to current problems in domestication research, and demonstrate the ways in which simulation modeling is beginning to reshape our understanding of the domestication process.

Multilevel and kin selection in a connected world.

Wild et al. argue that the evolution of reduced virulence can be understood from the perspective of inclusive fitness, obviating the need to evoke group selection as a contributing causal factor. Although they acknowledge the mathematical equivalence of the inclusive fitness and multilevel selection approaches, they conclude that reduced virulence can be viewed entirely as an individual-level adaptation by the parasite. Here we show that their model is a well-known special case of the more general theory of multilevel selection, and that the cause of reduced virulence resides in the opposition of two processes: within-group and among-group selection. This distinction is important in light of the current controversy among evolutionary biologists in which some continue to affirm that natural selection centres only and always at the level of the individual organism or gene, despite mathematical demonstrations that evolutionary dynamics must be described by selection at various levels in the hierarchy of biological organization.

Cultural group selection follows Darwin's classic syllogism for the operation of selection.

The main objective of our target article was to sketch the empirical case for the importance of selection at the level of groups on cultural variation. Such variation is massive in humans, but modest or absent in other species. Group selection processes acting on this variation is a framework for developing explanations of the unusual level of cooperation between non-relatives found in our species. Our case for cultural group selection (CGS) followed Darwin's classic syllogism regarding natural selection: If variation exists at the level of groups, if this variation is heritable, and if it plays a role in the success or failure of competing groups, then selection will operate at the level of groups. We outlined the relevant domains where such evidence can be sought and characterized the main conclusions of work in those domains. Most commentators agree that CGS plays some role in human evolution, although some were considerably more skeptical. Some contributed additional empirical cases. Some raised issues of the scope of CGS explanations versus competing ones.

Cultural group selection plays an essential role in explaining human cooperation: A sketch of the evidence.

Human cooperation is highly unusual. We live in large groups composed mostly of non-relatives. Evolutionists have proposed a number of explanations for this pattern, including cultural group selection and extensions of more general processes such as reciprocity, kin selection, and multi-level selection acting on genes. Evolutionary processes are consilient; they affect several different empirical domains, such as patterns of behavior and the proximal drivers of that behavior. In this target article, we sketch the evidence from five domains that bear on the explanatory adequacy of cultural group selection and competing hypotheses to explain human cooperation. Does cultural transmission constitute an inheritance system that can evolve in a Darwinian fashion? Are the norms that underpin institutions among the cultural traits so transmitted? Do we observe sufficient variation at the level of groups of considerable size for group selection to be a plausible process? Do human groups compete, and do success and failure in competition depend upon cultural variation? Do we observe adaptations for cooperation in humans that most plausibly arose by cultural group selection? If the answer to one of these questions is "no," then we must look to other hypotheses. We present evidence, including quantitative evidence, that the answer to all of the questions is "yes" and argue that we must take the cultural group selection hypothesis seriously. If culturally transmitted systems of rules (institutions) that limit individual deviance organize cooperation in human societies, then it is not clear that any extant alternative to cultural group selection can be a complete explanation.

Individual consistency and flexibility in human social information use.

Copying others appears to be a cost-effective way of obtaining adaptive information, particularly when flexibly employed. However, adult humans differ considerably in their propensity to use information from others, even when this 'social information' is beneficial, raising the possibility that stable individual differences constrain flexibility in social information use. We used two dissimilar decision-making computer games to investigate whether individuals flexibly adjusted their use of social information to current conditions or whether they valued social information similarly in both games. Participants also completed established personality questionnaires. We found that participants demonstrated considerable flexibility, adjusting social information use to current conditions. In particular, individuals employed a 'copy-when-uncertain' social learning strategy, supporting a core, but untested, assumption of influential theoretical models of cultural transmission. Moreover, participants adjusted the amount invested in their decision based on the perceived reliability of personally gathered information combined with the available social information. However, despite this strategic flexibility, participants also exhibited consistent individual differences in their propensities to use and value social information. Moreover, individuals who favoured social information self-reported as more collectivist than others. We discuss the implications of our results for social information use and cultural transmission.

The cultural niche: why social learning is essential for human adaptation.

In the last 60,000 y humans have expanded across the globe and now occupy a wider range than any other terrestrial species. Our ability to successfully adapt to such a diverse range of habitats is often explained in terms of our cognitive ability. Humans have relatively bigger brains and more computing power than other animals, and this allows us to figure out how to live in a wide range of environments. Here we argue that humans may be smarter than other creatures, but none of us is nearly smart enough to acquire all of the information necessary to survive in any single habitat. In even the simplest foraging societies, people depend on a vast array of tools, detailed bodies of local knowledge, and complex social arrangements and often do not understand why these tools, beliefs, and behaviors are adaptive. We owe our success to our uniquely developed ability to learn from others. This capacity enables humans to gradually accumulate information across generations and develop well-adapted tools, beliefs, and practices that are too complex for any single individual to invent during their lifetime.

Many important group-level traits are institutions.

Smaldino makes a solid contribution to the literature on the evolution of human social organization by pointing out that group-level-traits (GLTs) often emerge from the interactions of group members in such a way that their effects are not easily partitioned into individual selection. However, we argue that he too readily dismisses institutional analysis as a tool for understanding these traits.

Colloquium paper: gene-culture coevolution in the age of genomics.

The use of socially learned information (culture) is central to human adaptations. We investigate the hypothesis that the process of cultural evolution has played an active, leading role in the evolution of genes. Culture normally evolves more rapidly than genes, creating novel environments that expose genes to new selective pressures. Many human genes that have been shown to be under recent or current selection are changing as a result of new environments created by cultural innovations. Some changed in response to the development of agricultural subsistence systems in the Early and Middle Holocene. Alleles coding for adaptations to diets rich in plant starch (e.g., amylase copy number) and to epidemic diseases evolved as human populations expanded (e.g., sickle cell and G6PD deficiency alleles that provide protection against malaria). Large-scale scans using patterns of linkage disequilibrium to detect recent selection suggest that many more genes evolved in response to agriculture. Genetic change in response to the novel social environment of contemporary modern societies is also likely to be occurring. The functional effects of most of the alleles under selection during the last 10,000 years are currently unknown. Also unknown is the role of paleoenvironmental change in regulating the tempo of hominin evolution. Although the full extent of culture-driven gene-culture coevolution is thus far unknown for the deeper history of the human lineage, theory and some evidence suggest that such effects were profound. Genomic methods promise to have a major impact on our understanding of gene-culture coevolution over the span of hominin evolutionary history.

Culture rather than genes provides greater scope for the evolution of large-scale human prosociality.

Whether competition among large groups played an important role in human social evolution is dependent on how variation, whether cultural or genetic, is maintained between groups. Comparisons between genetic and cultural differentiation between neighboring groups show how natural selection on large groups is more plausible on cultural rather than genetic variation.

The punishment that sustains cooperation is often coordinated and costly.

Experiments are not models of cooperation; instead, they demonstrate the presence of the ethical and other-regarding predispositions that often motivate cooperation and the punishment of free-riders. Experimental behavior predicts subjects' cooperation in the field. Ethnographic studies in small-scale societies without formal coercive institutions demonstrate that disciplining defectors is both essential to cooperation and often costly to the punisher.

Authority matters: propaganda and the coevolution of behaviour and attitudes.

Human decision-making is controlled by various factors including material cost-benefit considerations, values and beliefs, social influences, cognitive factors and errors. Among social influences, those by external authorities (e.g. educational, cultural, religious, political, administrative, etc.) are particularly important owing to their potential reach and power. To better understand the effects of 'soft' power of authorities we develop a unifying theoretical framework integrating material, cognitive and social forces controlling the joint dynamics of individual actions and beliefs. We apply our approach to three different phenomena: evolution of food sharing in small-scale societies, participation in political protests and effects of priming social identity in behavioural experiments. For each of these applications, we show that our approach leads to different (or simpler) explanations of human behaviour than alternatives. We highlight the type of measurements which can be helpful in developing practical applications of our approach. We identify and explicitly characterise the degree of mismatch between individual actions and attitudes. We assert that the effects of external authorities, of changing beliefs and of differences between people must be studied empirically, included in mathematical models, and accounted for when developing different policies aiming to modify or sustain human behaviour.

Modern theories of human evolution foreshadowed by Darwin's Descent of Man.

Charles Darwin's The Descent of Man, published 150 years ago, laid the grounds for scientific studies into human origins and evolution. Three of his insights have been reinforced by modern science. The first is that we share many characteristics (genetic, developmental, physiological, morphological, cognitive, and psychological) with our closest relatives, the anthropoid apes. The second is that humans have a talent for high-level cooperation reinforced by morality and social norms. The third is that we have greatly expanded the social learning capacity that we see already in other primates. Darwin's emphasis on the role of culture deserves special attention because during an increasingly unstable Pleistocene environment, cultural accumulation allowed changes in life history; increased cognition; and the appearance of language, social norms, and institutions.