Investigating the effects of social information on spite in an online game.

While humans are highly cooperative, they can also behave spitefully. Yet spite remains understudied. Spite can be normatively driven and while previous experiments have found some evidence that cooperation and punishment may spread via social learning, no experiments have considered the social transmission of spiteful behaviour. Here we present an online experiment where, following an opportunity to earn wealth, we asked participants to choose an action towards an anonymous partner across a full spectrum of social behaviour, from spite to altruism. In accordance with cultural evolutionary theory, participants were presented with social information that varied in source and content. Across six conditions, we informed participants that either the majority or the highest earner had chosen to behave spitefully, neutrally or altruistically. We found an overall tendency towards altruism, but at lower levels among those exposed to spite compared with altruism. We found no difference between social information that came from the majority or the highest earner. Exploratory analysis revealed that participants' earnings negatively correlated with altruistic behaviour. Our results contrast with previous literature that report high rates of spite in experimental samples and a greater propensity for individuals to copy successful individuals over the majority.

Cultural niche construction with application to fertility control: A model for education and social transmission of contraceptive use.

The evolution of a cultural trait may be affected by niche construction, or changes in the selective environment of that trait due to the inheritance of other cultural traits that make up a cultural background. This study investigates the evolution of a cultural trait, such as the acceptance of the idea of contraception, that is both vertically and horizontally transmitted within a homogeneous social network. Individuals may conform to the norm, and adopters of the trait have fewer progeny than others. In addition, adoption of this trait is affected by a vertically transmitted aspect of the cultural background, such as the preference for high or low levels of education. Our model shows that such cultural niche construction can facilitate the spread of traits with low Darwinian fitness while providing an environment that counteracts conformity to norms. In addition, niche construction can facilitate the 'demographic transition' by making reduced fertility socially accepted.

Unknotting the interactive effects of learning processes on cultural evolutionary dynamics.

Forms of non-random copying error provide sources of inherited variation yet their effects on cultural evolutionary dynamics are poorly understood. Focusing on variation in granny and reef knot forms, we present a mathematical model that specifies how these variant frequencies are affected by non-linear interactions between copying fidelity, mirroring, handedness and repetition biases. Experiments on adult humans allowed these effects to be estimated using approximate Bayesian computation and the model is iterated to explain the prevalence of granny over reef knots in the wild. Our study system also serves to show conditions under which copying fidelity drives heterogeneity in cultural variants at equilibrium, and that interaction between unbiased forms of copying error can skew cultural variation.

Style of pictorial representation is shaped by intergroup contact.

Pictorial representation is a key human behaviour. Cultures around the world have made images to convey information about living kinds, objects and ideas for at least 75,000 years, in forms as diverse as cave paintings, religious icons and emojis. However, styles of pictorial representation vary greatly between cultures and historical periods. In particular, they can differ in figurativeness, i.e. varying from detailed depictions of subjects to stylised abstract forms. Here we show that pictorial styles can be shaped by intergroup contact. We use data from experimental microsocieties to show that drawings produced by groups in contact tended to become more figurative and transparent to outsiders, whereas in isolated groups drawings tended to become abstract and opaque. These results indicate that intergroup contact is likely to be an important factor in the cultural evolution of pictorial representation, because the need to communicate with outsiders ensures that some figurativeness is retained over time. We discuss the implications of this finding for understanding the history and anthropology of art, and the parallels with sociolinguistics and language evolution.

Multidisciplinary exhibit design in a Science Centre: a participatory action research approach.

In this paper we highlight the issues and opportunities of a participatory action research (PAR) and co-design project, currently being undertaken as engaged research between academics at Durham University and practitioners at the UK's International Centre for Life in Newcastle-Upon-Tyne (CfL; see creativescienceatlife.com for more information and developments). The focus is on the use of PAR to enable university researchers and Science Centre professionals to co-design Informal Science Learning exhibits that enhance creativity and innovation in young people. We define the principles of PAR and explore reasons for adopting the approach. An account is provided of the iterative co-design and piloting of a novel exhibit within a new exhibition space at the CfL. Reflections collated independently by the practitioners and the academics involved highlighting the development of ideas and insights over the course of the PAR process. We discuss how PAR enabled effective engagement with and creation of enriched knowledge, and innovation, in both the academy and science-learning professionals. The added value of PAR and co-production to our project aligns with current calls for a redefining of how societal impact of academic research is considered.

Testing differential use of payoff-biased social learning strategies in children and chimpanzees.

Various non-human animal species have been shown to exhibit behavioural traditions. Importantly, this research has been guided by what we know of human culture, and the question of whether animal cultures may be homologous or analogous to our own culture. In this paper, we assess whether models of human cultural transmission are relevant to understanding biological fundamentals by investigating whether accounts of human payoff-biased social learning are relevant to chimpanzees (Pan troglodytes). We submitted 4- and 5-year-old children (N = 90) and captive chimpanzees (N = 69) to a token-reward exchange task. The results revealed different forms of payoff-biased learning across species and contexts. Specifically, following personal and social exposure to different tokens, children's exchange behaviour was consistent with proportional imitation, where choice is affected by both prior personally acquired and socially demonstrated token-reward information. However, when the socially derived information regarding token value was novel, children's behaviour was consistent with proportional observation; paying attention to socially derived information and ignoring their prior personal experience. By contrast, chimpanzees' token choice was governed by their own prior experience only, with no effect of social demonstration on token choice, conforming to proportional reservation. We also find evidence for individual- and group-level differences in behaviour in both species. Despite the difference in payoff strategies used, both chimpanzees and children adopted beneficial traits when available. However, the strategies of the children are expected to be the most beneficial in promoting flexible behaviour by enabling existing behaviours to be updated or replaced with new and often superior ones.

Fire in the Plio-Pleistocene: the functions of hominin fire use, and the mechanistic, developmental and evolutionary consequences.

Fire is a powerful natural force that can change landscapes extremely quickly. Hominins have harnessed this resource for their own purposes, with mechanistic and developmental physiological consequences. In addition, the use of fire has niche constructive effects, altering selective environments for genetic and cultural evolution. We review the record for hominin fire use in the Plio-Pleistocene, before considering the various functions for its use, and the resultant mechanistic and developmental consequences. We also adopt the niche construction framework to consider how the use of fire can modify selective environments, and thus have evolutionary consequences at genetic and cultural levels. The light that fire produces may influence photoperiodicity and alter hormonally-controlled bodily rhythms. Fire used for cooking could have extended the range of foods hominins were able to consume, and reduced digestion costs. This may have contributed to the expansion of the hominin brain and facial anatomy, influenced by a higher quality cooked diet. Fire may also have allowed dispersal into northern areas with much cooler climates than the hominin African origin, posing novel problems that affected diet and social behaviour.

An SIS model for cultural trait transmission with conformity bias.

Epidemiological models have been applied to human health-related behaviors that are affected by social interaction. Typically these models have not considered conformity bias, that is, the exaggerated propensity to adopt commonly observed behaviors or opinions, or content biases, where the content of the learned trait affects the probability of adoption. Here we consider an interaction of these two effects, presenting an SIS-type model for the spread and persistence of a behavior which is transmitted via social learning. Uptake is controlled by a nonlinear dependence on the proportion of individuals demonstrating the behavior in a population. Three equilibrium solutions are found, their linear stability is analyzed and the results are compared with a model for unbiased social learning. Our analysis focuses on the effects of the strength of conformity bias and the effects of content biases which alter a conformity threshold frequency of the behavior, above which there is an exaggerated propensity for adoption. The strength of the conformity bias is found to qualitatively alter the predictions regarding whether the trait becomes endemic within the population and the proportion of individuals who display the trait when it is endemic. As the conformity strength increases, the number of feasible equilibrium solutions increases from two to three, leading to a situation where the stable equilibrium attained is dependent upon the initial state. Varying the conformity threshold frequency directionally alters the behavior invasion threshold. Finally we discuss the possible application of this model to binge drinking behavior.

Target article with commentaries: developmental niche construction.

Niche construction is the modification of components of the environment through an organism's activities. Humans modify their environments mainly through ontogenetic and cultural processes, and it is this reliance on learning, plasticity and culture that lends human niche construction a special potency. In this paper we aim to facilitate discussion between researchers interested in niche construction and those interested in human cognitive development by highlighting some of the related processes. We discuss the transmission of culturally relevant information, how the human mind is a symbol-generating and artefact-devising system, and how these processes are bi-directional, with infants and children both being directed, and directing, their own development. We reflect on these in the light of four approaches: natural pedagogy, activity theory, distributed cognition and situated learning. Throughout, we highlight pertinent examples in non-humans that parallel or further explicate the processes discussed. Finally we offer three future directions; two involving the use of new techniques in the realms of neuroscience and modelling, and the third suggesting exploration of changes in the effects of niche construction across the lifespan.

Human niche construction in interdisciplinary focus.

Niche construction is an endogenous causal process in evolution, reciprocal to the causal process of natural selection. It works by adding ecological inheritance, comprising the inheritance of natural selection pressures previously modified by niche construction, to genetic inheritance in evolution. Human niche construction modifies selection pressures in environments in ways that affect both human evolution, and the evolution of other species. Human ecological inheritance is exceptionally potent because it includes the social transmission and inheritance of cultural knowledge, and material culture. Human genetic inheritance in combination with human cultural inheritance thus provides a basis for gene-culture coevolution, and multivariate dynamics in cultural evolution. Niche construction theory potentially integrates the biological and social aspects of the human sciences. We elaborate on these processes, and provide brief introductions to each of the papers published in this theme issue.

Do non-human primates synchronise their menstrual cycles? A test in mandrills.

The concept of female cycle (or estrous) synchrony has enduring popular appeal. However, critical reviews of estrous synchrony studies in both humans and non-humans have found that synchrony has not been demonstrated convincingly, due to methodological artifacts and statistical problems. Studies of this phenomenon in animals living under naturalistic conditions are rare. We used long-term records of the timing of the female menstrual cycle in a semi-free-ranging population of mandrills, together with a randomisation procedure, to test hypotheses relating to cycle synchrony in a naturally reproducing primate. We found evidence of significant synchrony of the peri-ovulatory period in only one of 10 group-years - the year in which the largest number of cycles was recorded, both overall and per female. However, this result was no longer significant when we corrected for multiple tests of the same hypothesis. This suggests that mandrills in our study population do not synchronise their cycles, possibly because they usually conceive so quickly that they do not have the opportunity to synchronise. We also tested whether females in the same matriline, which associate with one another more than other females, cycle significantly more closely together in time than unrelated females, finding that they did so in 2 of 10 group-years, but that they were significantly less likely to match their cycles in another group-year. Across 32 matriline-years, patterns of synchrony within individual matrilines (female lineages) never fell outside the distribution based on chance. Thus we found little support for the pheromonal hypothesis for cycle synchrony, which predicts that females that associate with one another should be more likely to cycle together. Overall, our findings are in line with other studies that suggest that cycle synchrony does not occur in non-human primates.

Evidence for social learning in wild lemurs (Lemur catta).

Interest in social learning has been fueled by claims of culture in wild animals. These remain controversial because alternative explanations to social learning, such as asocial learning or ecological differences, remain difficult to refute. Compared with laboratory-based research, the study of social learning in natural contexts is in its infancy. Here, for the first time, we apply two new statistical methods, option-bias analysis and network-based diffusion analysis, to data from the wild, complemented by standard inferential statistics. Contrary to common thought regarding the cognitive abilities of prosimian primates, our evidence is consistent with social learning within subgroups in the ring-tailed lemur (Lemur catta), supporting the theory of directed social learning (Coussi-Korbel & Fragaszy, 1995). We also caution that, as the toolbox for capturing social learning in natural contexts grows, care is required in ensuring that the methods employed are appropriate-in particular, regarding social dynamics among study subjects. Supplemental materials for this article may be downloaded from http://lb.psychonomic-journals.org/content/supplemental.

The effect of task structure on diffusion dynamics: Implications for diffusion curve and network-based analyses.

Theoretical analyses within the broad field of social learning research give mixed conclusions on whether the shape of a diffusion curve can be used to infer that a learned trait increases through social or asocial learning. Here we explore how factors such as task structure (e.g., multiple-step tasks), task abandonment, subgoal learning, and neophobia affect the shape of the diffusion curve for both asocially learned and socially learned behavior. We demonstrate that, whereas social learning increases the likelihood of S-shaped curves, sigmoidal patterns can be generated by entirely asocial processes, and cannot be reliably interpreted as indicators of social learning. Our findings reinforce the view that diffusion curve analysis is not a reliable way of detecting social transmission. We also draw attention to the fact that task structure can similarly confound interpretation of network-based diffusion analyses, and suggest resolutions to this problem. Supplemental materials for this article may be downloaded from http://lb.psychonomic-journals.org/content/supplemental.

Identifying social learning in animal populations: a new 'option-bias' method.

BACKGROUND: Studies of natural animal populations reveal widespread evidence for the diffusion of novel behaviour patterns, and for intra- and inter-population variation in behaviour. However, claims that these are manifestations of animal 'culture' remain controversial because alternative explanations to social learning remain difficult to refute. This inability to identify social learning in social settings has also contributed to the failure to test evolutionary hypotheses concerning the social learning strategies that animals deploy. METHODOLOGY/PRINCIPAL FINDINGS: We present a solution to this problem, in the form of a new means of identifying social learning in animal populations. The method is based on the well-established premise of social learning research, that--when ecological and genetic differences are accounted for--social learning will generate greater homogeneity in behaviour between animals than expected in its absence. Our procedure compares the observed level of homogeneity to a sampling distribution generated utilizing randomization and other procedures, allowing claims of social learning to be evaluated according to consensual standards. We illustrate the method on data from groups of monkeys provided with novel two-option extractive foraging tasks, demonstrating that social learning can indeed be distinguished from unlearned processes and a social learning, and revealing that the monkeys only employed social learning for the more difficult tasks. The method is further validated against published datasets and through simulation, and exhibits higher statistical power than conventional inferential statistics. CONCLUSIONS/SIGNIFICANCE: The method is potentially a significant technological development, which could prove of considerable value in assessing the validity of claims for culturally transmitted behaviour in animal groups. It will also be of value in enabling investigation of the social learning strategies deployed in captive and natural animal populations.

The evolution of social learning rules: payoff-biased and frequency-dependent biased transmission.

Humans and other animals do not use social learning indiscriminately, rather, natural selection has favoured the evolution of social learning rules that make selective use of social learning to acquire relevant information in a changing environment. We present a gene-culture coevolutionary analysis of a small selection of such rules (unbiased social learning, payoff-biased social learning and frequency-dependent biased social learning, including conformism and anti-conformism) in a population of asocial learners where the environment is subject to a constant probability of change to a novel state. We define conditions under which each rule evolves to a genetically polymorphic equilibrium. We find that payoff-biased social learning may evolve under high levels of environmental variation if the fitness benefit associated with the acquired behaviour is either high or low but not of intermediate value. In contrast, both conformist and anti-conformist biases can become fixed when environment variation is low, whereupon the mean fitness in the population is higher than for a population of asocial learners. Our examination of the population dynamics reveals stable limit cycles under conformist and anti-conformist biases and some highly complex dynamics including chaos. Anti-conformists can out-compete conformists when conditions favour a low equilibrium frequency of the learned behaviour. We conclude that evolution, punctuated by the repeated successful invasion of different social learning rules, should continuously favour a reduction in the equilibrium frequency of asocial learning, and propose that, among competing social learning rules, the dominant rule will be the one that can persist with the lowest frequency of asocial learning.

From traditional medicine to witchcraft: why medical treatments are not always efficacious.

Complementary medicines, traditional remedies and home cures for medical ailments are used extensively world-wide, representing more than US$60 billion sales in the global market. With serious doubts about the efficacy and safety of many treatments, the industry remains steeped in controversy. Little is known about factors affecting the prevalence of efficacious and non-efficacious self-medicative treatments. Here we develop mathematical models which reveal that the most efficacious treatments are not necessarily those most likely to spread. Indeed, purely superstitious remedies, or even maladaptive practices, spread more readily than efficacious treatments under specified circumstances. Low-efficacy practices sometimes spread because their very ineffectiveness results in longer, more salient demonstration and a larger number of converts, which more than compensates for greater rates of abandonment. These models also illuminate a broader range of phenomena, including the spread of innovations, medical treatment of animals, foraging behaviour, and self-medication in non-human primates.

Conceptual Barriers to Progress Within Evolutionary Biology.

In spite of its success, Neo-Darwinism is faced with major conceptual barriers to further progress, deriving directly from its metaphysical foundations. Most importantly, neo-Darwinism fails to recognize a fundamental cause of evolutionary change, "niche construction". This failure restricts the generality of evolutionary theory, and introduces inaccuracies. It also hinders the integration of evolutionary biology with neighbouring disciplines, including ecosystem ecology, developmental biology, and the human sciences. Ecology is forced to become a divided discipline, developmental biology is stubbornly difficult to reconcile with evolutionary theory, and the majority of biologists and social scientists are still unhappy with evolutionary accounts of human behaviour. The incorporation of niche construction as both a cause and a product of evolution removes these disciplinary boundaries while greatly generalizing the explanatory power of evolutionary theory.

Quantifying and modelling social learning processes in monkey populations

La existencia de supuestas tradiciones en las poblaciones animales sigue sin resolverse enausencia de métodos que permitan aislar los mecanismos de transmisión social en losgrupos sociales. Aquí tratamos este problema mediante la cuantificación del efecto quejugaron dos procesos de aprendizaje social, el realce del estímulo y el aprendizajeobservacional, en la adopción de un comportamiento novedoso de recolección en gruposde monos de la familia Callitrichidae. Simulamos el efecto de estos procesos en unmodelo de cara a explicar la extensión de un comportamiento novedoso y para seleccionarque parámetros del modelo proporcionaban el mejor ajuste a los datos del mono.Nuestro análisis revela evidencias para los procesos asociales y el realce estimular, perono para aprendizaje observacional. Las latencias empleadas en la solución de las tareascovarió con la fuerza de los procesos asociales, pero no con la de los procesos sociales.Más aún, el modelo solo requirió los parámetros de los factores no sociales para ajustarlos datos. El modelo y los datos de difusión mostraron ambos curvas desaceleradas dedifusión. Finalmente, discutimos la relación entre los procesos asociales y sociales y ladinámica de la difusión cultural(AU)

Putative traditions in animal populations remain unsubstantiated in the absence of methodsto isolate the mechanisms of social transmission in social groups. Here we address thisproblem by quantifying the effect of two social learning processes, namely stimulusenhancement and observational learning, on the adoption of a novel extractive foragingbehavior in groups of callitrichid monkeys. We simulate the effect of these processes ina model for the spread of a novel behaviour and also select which model-parametersprovide the best fit to the monkey data. Our analysis reveals evidence for asocial processesand stimulus enhancement but not observational learning. The latency to solve the tasksco-varied with the strength of the asocial, but not social, processes. Further, only asocialparameters were required for the model to fit the data. Both model and monkey diffusiondata exhibited deceleratory diffusion curves. We discuss the relationship between both theasocial and social processes and the diffusion dynamics(AU)

Cultural coevolution of norm adoption and enforcement when punishers are rewarded or non-punishers are punished.

A number of studies have shown that social norms can be maintained at a high frequency when norm-violators are punished. However, there remains the problem of how norm-adopters and punishers coevolve within a single group. We develop a recursive system to examine the coevolution of norm-adopters and punishers where the viability of punishers is enhanced by one of two "metanorms": (1) Norm-observers reward punishers for punishing norm-violators (Reward Model); (2) Punishers punish non-punishers (Punishment Model). Both models generate a bistable system and each is characterized in phenotype frequency space by a distinct region of attraction to the equilibrium consisting of only norm-adopting punishers. Using a Monte Carlo simulation, we find that cultural drift may allow norm-adopters and punishers to coevolve from invasion into this region of attraction, resulting in their fixation. This coevolution typically occurs across a wider range of conditions under the reward- than the punishment-based metanorm. We also show that, under appropriate conditions, a large negative statistical association between the two traits may evolve only under the Reward Model. Furthermore, for each metanorm, a population of norm-adopters who always observe the norm can be locally stable over a continuum of punishment frequencies.

Cultural niche construction in a metapopulation.

Cultural niche construction is the process by which certain evolving cultural traits form a cultural niche that affects the evolution of other genetic and cultural traits [Laland, K., et al., 2001. Cultural niche construction and human evolution. J. Evol. Biol. 14, 22-33; Ihara, Y., Feldman, M., 2004. Cultural niche construction and the evolution of small family size. Theor. Popul. Biol. 65, 105-111]. In this study we focus on cultural niche construction in a metapopulation (a population of populations), where the frequency of one cultural trait (e.g. the level of education) determines the transmission rate of a second trait (e.g. the adoption of fertility reduction preferences) within and between populations. We formulate the Metapopulation Cultural Niche Construction (MPCNC) model by defining the cultural niche induced by the first trait as the construction of a social interaction network on which the second trait may percolate. Analysis of the model reveals dynamics that are markedly different from those observed in a single population, allowing, for example, different (or even opposing) dynamics in each population. In particular, this model can account for the puzzling phenomenon reported in previous studies [Bongaarts, J., Watkins, S., 1996. Social interactions and contemporary fertility transitions. Popul. Dev. Rev. 22 (4), 639-682] that the onset of the demographic transition in different countries occurred at ever lower levels of development.