The evolution of conformist social learning can cause population collapse in realistically variable environments

Why do societies collapse? We use an individual-based evolutionary model to show that, in environmental conditions dominated by low-frequency variation (“red noise”), extirpation may be an outcome of the evolution of cultural capacity. Previous analytical models predicted an equilibrium between individual learners and social learners, or a contingent strategy in which individuals learn socially or individually depending on the circumstances. However, in red noise environments, whose main signature is that variation is concentrated in relatively large, relatively rare excursions, individual learning may be selected from the population. If the social learning system comes to lack sufficient individual learning or cognitively costly adaptive biases, behavior ceases tracking environmental variation. Then, when the environment does change, fitness declines and the population may collapse or even be extirpated. The modeled scenario broadly fits some human population collapses and might also explain nonhuman extirpations. Varying model parameters showed that the fixation of social learning is less likely when individual learning is less costly, when the environment is less red or more variable, with larger population sizes, and when learning is not conformist or is from parents rather than from the general population. Once social learning is fixed, extirpation is likely except when social learning is biased towards successful models. Thus, the risk of population collapse may be reduced by promoting individual learning and innovation over cultural conformity, or by preferential selection of relatively fit individuals as models for social learning.     

The evolution of conformist transmission in social learning when the environment changes periodically

Conformity is often observed in human social learning. Social learners preferentially imitate the majority or most common behavior in many situations, though the strength of conformity varies with the situation. Why has such a psychological tendency evolved? I investigate this problem by extending a standard model of social learning evolution with infinite environmental states (Feldman, M.W., Aoki, K., Kumm, J., 1996. Individual versus social learning: evolutionary analysis in a fluctuating environment. Anthropol. Sci. 104, 209-231) to include conformity bias. I mainly focus on the relationship between the strength of conformity bias that evolves and environmental stability, which is one of the most important factors in the evolution of social learning. Using the evolutionarily stable strategy (ESS) approach, I show that conformity always evolves when environmental stability and the cost of adopting a wrong behavior are small, though environmental stability and the cost of individual learning both negatively affect the strength of conformity.     

Interdependent utilities: how social ranking affects choice behavior

Organization in hierarchical dominance structures is prevalent in animal societies, so a strong preference for higher positions in social ranking is likely to be an important motivation of human social and economic behavior. This preference is also likely to influence the way in which we evaluate our outcome and the outcome of others, and finally the way we choose. In our experiment participants choose among lotteries with different levels of risk, and can observe the choice that others have made. Results show that the relative weight of gains and losses is the opposite in the private and social domain. For private outcomes, experience and anticipation of losses loom larger than gains, whereas in the social domain, gains loom larger than losses, as indexed by subjective emotional evaluations and physiological responses. We propose a theoretical model (interdependent utilities), predicting the implication of this effect for choice behavior. The relatively larger weight assigned to social gains strongly affects choices, inducing complementary behavior: faced with a weaker competitor, participants adopt a more risky and dominant behavior.     

Estimating Impacts of Climate Change Policy on Land Use: An Agent-Based Modelling Approach

Agriculture is important to New Zealand’s economy. Like other primary producers, New Zealand strives to increase agricultural output while maintaining environmental integrity. Utilising modelling to explore the economic, environmental and land use impacts of policy is critical to understand the likely effects on the sector. Key deficiencies within existing land use and land cover change models are the lack of heterogeneity in farmers and their behaviour, the role that social networks play in information transfer, and the abstraction of the global and regional economic aspects within local-scale approaches. To resolve these issues we developed the Agent-based Rural Land Use New Zealand model. The model utilises a partial equilibrium economic model and an agent-based decision-making framework to explore how the cumulative effects of individual farmer’s decisions affect farm conversion and the resulting land use at a catchment scale. The model is intended to assist in the development of policy to shape agricultural land use intensification in New Zealand. We illustrate the model, by modelling the impact of a greenhouse gas price on farm-level land use, net revenue, and environmental indicators such as nutrient losses and soil erosion for key enterprises in the Hurunui and Waiau catchments of North Canterbury in New Zealand. Key results from the model show that farm net revenue is estimated to increase over time regardless of the greenhouse gas price. Net greenhouse gas emissions are estimated to decline over time, even under a no GHG price baseline, due to an expansion of forestry on low productivity land. Higher GHG prices provide a greater net reduction of emissions. While social and geographic network effects have minimal impact on net revenue and environmental outputs for the catchment, they do have an effect on the spatial arrangement of land use and in particular the clustering of enterprises.     

Applying evolutionary models to the laboratory study of social learning

Cultural evolution is driven, in part, by the strategies that individuals employ to acquire behavior from others. These strategies themselves are partly products of natural selection, making the study of social learning an inherently Darwinian project. Formal models of the evolution of social learning suggest that reliance on social learning should increase with task difficulty and decrease with the probability of environmental change. These models also make predictions about how individuals integrate information from multiple peers. We present the results of microsociety experiments designed to evaluate these predictions. The first experiment measures baseline individual learning strategy in a two-armed bandit environment with variation in task difficulty and temporal fluctuation in the payoffs of the options. Our second experiment addresses how people in the same environment use minimal social information from a single peer. Our third experiment expands on the second by allowing access to the behavior of several other individuals, permitting frequency-dependent strategies like conformity. In each of these experiments, we vary task difficulty and environmental fluctuation. We present several candidate strategies and compute the expected payoffs to each in our experimental environment. We then fit to the data the different models of the use of social information and identify the best-fitting model via model comparison techniques. We find substantial evidence of both conformist and nonconformist social learning and compare our results to theoretical expectations.     

Learning,productivity, and noise: an experimental study of cultural transmission on the Bolivian Altiplano

The theory of cultural transmission distinguishes between biased and unbiased social learning. Biases simply mean that social learning is not completely random. The distinction is critical because biases produce effects at the aggregate level that then feed back to influence individual behavior. This study presents an economic experiment designed specifically to see if players use social information in a biased way. The experiment was conducted among a group of subsistence pastoralists in southern Bolivia. Treatments were designed to test for two widely discussed forms of biased social learning: a tendency to imitate success and a tendency to follow the majority. The analysis, based primarily on fitting specific evolutionary models to the data using maximum likelihood, found neither a clear tendency to imitate success nor conformity. Players instead seemed to rely largely on private feedback about their own personal histories of choices and payoffs. Nonetheless, improved performance in one treatment provides evidence for some important but currently unspecified social effect. Given existing experimental work on cultural transmission from other societies, the current study suggests that social learning is potentially conditional and culturally specific.     

The evolutionary roots of our environmental problems: toward a Darwinian ecology

It is widely acknowledged that we need to stabilize population growth and reduce our environmental impact; however, there is little consensus about how we might achieve these changes. Here I show how evolutionary analyses of human behavior provide important, though generally ignored, insights into our environmental problems. First, I review increasing evidence that Homo sapiens has a long history of causing ecological problems. This means that, contrary to popular belief, our species' capacity for ecological destruction is not simply due to "Western" culture. Second, I provide an overview of how evolutionary research can help to understand why humans are ecologically destructive, including the reasons why people often overpopulate, overconsume, exhaust common-pool resources, discount the future, and respond maladaptively to modern environmental hazards. Evolutionary approaches not only explain our darker sides, they also provide insights into why people cherish plants and animals and often support environmental and conservation efforts (e.g., Wilson's "biophilia hypothesis"). Third, I show how evolutionary analyses of human behavior offer practical implications for environmental policy, education, and activism. I suggest that education is necessary but insufficient because people also need incentives. Individual incentives are likely to be the most effective, but these include much more than narrow economic interests (e.g., they include one's reputation in society). Moralizing and other forms of social pressure used by environmentalists to bring about change appear to be effective, but this idea needs more research. Finally, I suggest that integrating evolutionary perspectives into the environmental sciences will help to break down the artificial barriers that continue to divide the biological and social sciences, which unfortunately obstruct our ability to understand ourselves and effectively address our environmental problems.     

Kin-selection,reciprocal altruism,and information sharing among Maine lobstermen

In this paper, the sharing of information about the location of lobsters among lobstermen in two Maine harbors is described. First, why the sharing of such information is likely to entail an economic loss for the transmitters is explained. Then, the extent to which the principles of kin-selection and reciprocal altruism can account for the sharing of information is determined. Many cases of information sharing in one of the harbors do not appear to be the type of kin-directed or reciprocal acts expected to be produced by kin-selection or reciprocal altruism as they are usually conceived. The behavior of these lobstermen may be the result of the advantages of maintaining a complex web of social relationships among them. Failure to appreciate the complexity of such relationships in some fishing communities is suggested to be a major shortcoming in the economic models previously used to explain information management among commercial fishermen. I conclude that a more complex model of reciprocal altruism is needed to account for the information sharing among this group of Maine lobstermen, and perhaps many other human social groups.     

On the Relative Importance of Haplo-Diploidy, Assortative Mating and Social Synergy on the Evolutionary Emergence of Social Behavior

Advances in multiagent simulation techniques make it possible to study more realistic dynamics of complex systems and allow evolutionary theories to be tested. Here I use simulations to assess the relative importance of reproductive systems (haplodiploidy vs. diploidy), mate selection (assortative mating vs. random mating) and social economics (pay-off matrices of evolutionary games) in the evolutionary dynamics leading to the emergence of social cooperation in the provision of parental care. The simulations confirm that haplo-diploid organisms and organisms mating assortatively have a higher probability for fixing alleles and require less favorable conditions for their fixation, than diploids or organisms mating randomly. The simulations showed that social behavior was most likely to emerge a) when the cost for parental investment was much lower than the benefits to the offspring, b) when cooperation improved synergistically the fitness of offspring compared to the corresponding egoistic behavior and c) when alleles coding for altruistic or social behavior could be rapidly fixed in the population, thanks to mechanisms such as haplo-diploidy and/or assortative mating. Cooperative social behavior always appeared if sociality conferred much higher fitness gains compared to non cooperative alternatives suggesting that the most important factors for the emergence and maintenance of social behavior are those based on energetic or efficiency considerations. The simulations, in congruence with the scant experimental evidence available, suggest that economic considerations rather than genetic ones are critical in explaining the emergence and maintenance of sociality.     

A theoretical investigation of the role of social transmission in evolution

This paper contains an investigation of the interaction between protocultural processes in animals, generated by social learning and the processes of biological evolution. It addresses the question of whether mechanisms of social learning and transmission can play an evolutionary role by allowing learned patterns of behavior to spread through animal populations, in the process changing the selection pressures acting on them. Simple models of social transmission and gene-meme coevolution are developed to investigate three hypotheses related to the role of social transmission in animal evolution. Simulations using the models suggest that social transmission would have to be particularly stable and be associated with estremely strong selection, if it were to result in the fixation of alleles. A more likely hypothesis is that social transmission might allow animals to respond adaptively to novelty in their environment, rendering a genetic response unnecessary, or only partially necessary. Socially transmitted traits appear to spread sufficiently rapidly, relative to changes in gene frequency, that it would be quite feasible for a socially transmitted response to an environmental change to occur, preempting a genetic response. Social transmission is probably more likely to slow down evolutionary rates than to speed them up through changing selection pressures. However, cultural and evolutionary processes are likely to interact in complex ways, and a “behavioral drive” effect cannot be ruled out.     

Social Learning in Noncolonial Insects?

Social-information use has generated great interest lately and has been shown to have important implications for the ecology and evolution of species. Learning about predators or predation risk from others may provide low-cost life-saving information and would be expected to have adaptive payoffs in any species where conspecifics are observable and behave differently under predation risk. Yet, social learning and social-information use in general have been largely restricted to vertebrates. Here, we show that crickets adapt their predator-avoidance behavior after having observed the behavior of knowledgeable others and maintain these behavioral changes lastingly after demonstrators are gone. These results point toward social learning, a contingency never shown before in noncolonial insects. We show that these long-lasting changes cannot instead be attributed to long re-emergence times, long-lasting effects of alarm pheromones, or residual odor cues. Our findings imply that social learning is likely much more phylogenetically widespread than currently acknowledged and that reliance on social information is determined by ecological rather than taxonomic constrains, and they question the generally held assumption that social learning is restricted to large-brained animals assumed to possess superior cognitive abilities.     

Learned social preference in zebrafish

How social aggregations arise and persist is central to our understanding of evolution, behavior, and psychology. When social groups arise within a species, evolutionary divergence and speciation can result. To understand this diversifying role of social behavior, we must examine the internal and external influences that lead to nonrandom assortment of phenotypes. Many fishes form aggregations called shoals that reduce predation risk while enhancing foraging and reproductive success. Thus, shoaling is adaptive, and signals that maintain shoals are likely to evolve under selection. Given the diversity of pigment patterns among Danio fishes, visual signals might be especially important in mediating social behaviors in this group. Our understanding of pigment pattern development in the zebrafish D. rerio allows integrative analyses of how molecular variation leads to morphological variation among individuals and how morphological variation influences social interactions. Here, we use the zebrafish pigment mutant nacre/mitfa to test roles for genetic and environmental determinants in the development of shoaling preference. We demonstrate that individuals discriminate between shoals having different pigment pattern phenotypes and that early experience determines shoaling preference. These results suggest a role for social learning in pigment pattern diversification in danios.     

Predicting resource utilization: the utility of optimal foraging models

Predicting how predation changes communities is an important challenge for ecologists. One view assumes that predictions can be made in terms of behavioural phenomena. Static optimality methods have been used to devise models of prey choice and are thought to have performed well. Two examples from the literature show that models of diet choice have either been applied without first testing them at the individual level, or have been tested in a way that does not fulfil all their assumptions. A discussion of the nature of mathematical models shows why it is dangerous to translate theories into natural language and how important it is to appreciate the limitations on the mathematical functions describing behaviour. A final section discusses behavioural phenomena such as satiation and learning, which are likely to limit the predictive capacities of static optimization models. It is likely that dynamic models are the way forward although they may be harder to apply at the ecological level.     

Language Learning and the Politics of Belonging: Sudanese Women Refugees Becoming and Being"American"

In this article, I explore the complicated relationship between ideologies of language and language learning, discourses of immigration and belonging, and the actual lived experiences of individual language learners. The analysis demonstrates how questions of educational access, economic stability, and social membership are all influenced by a range of social, political, and historical factors, particularly for recently arrived immigrants and refugees from war-torn African contexts.     

Social learning in cooperative dilemmas

Helping is a cornerstone of social organization and commonplace in human societies. A major challenge for the evolutionary sciences is to explain how cooperation is maintained in large populations with high levels of migration, conditions under which cooperators can be exploited by selfish individuals. Cultural group selection models posit that such large-scale cooperation evolves via selection acting on populations among which behavioural variation is maintained by the cultural transmission of cooperative norms. These models assume that individuals acquire cooperative strategies via social learning. This assumption remains empirically untested. Here, I test this by investigating whether individuals employ conformist or payoff-biased learning in public goods games conducted in 14 villages of a forager–horticulturist society, the Pahari Korwa of India. Individuals did not show a clear tendency to conform or to be payoff-biased and are highly variable in their use of social learning. This variation is partly explained by both individual and village characteristics. The tendency to conform decreases and to be payoff-biased increases as the value of the modal contribution increases. These findings suggest that the use of social learning in cooperative dilemmas is contingent on individuals'' circumstances and environments, and question the existence of stably transmitted cultural norms of cooperation.     

The utility of behavioral models and modules in molecular analyses of social behavior

It is extremely difficult to trace the causal pathway relating gene products or molecular pathways to the expression of behavior. This is especially true for social behavior, which being dependent on interactions and communication between individuals is even further removed from molecular-level events. In this review, we discuss how behavioral models can aid molecular analyses of social behavior. Various models of behavior exist, each of which suggest strategies to dissect complex behavior into simpler behavioral 'modules.' The resulting modules are easier to relate to neural processes and thus suggest hypotheses for neural and molecular function. Here we discuss how three different models of behavior have facilitated understanding the molecular bases of aspects of social behavior. We discuss the response threshold model and two different approaches to modeling motivation, the state space model and models of reinforcement and reward processing. The examples we have chosen illustrate how models can generate testable hypotheses for neural and molecular function and also how molecular analyses probe the validity of a model of behavior. We do not champion one model over another; rather, our examples illustrate how modeling and molecular analyses can be synergistic in exploring the molecular bases of social behavior.     

The influence of life history milestones and association networks on crop-raiding behavior in male African elephants

Factors that influence learning and the spread of behavior in wild animal populations are important for understanding species responses to changing environments and for species conservation. In populations of wildlife species that come into conflict with humans by raiding cultivated crops, simple models of exposure of individual animals to crops do not entirely explain the prevalence of crop raiding behavior. We investigated the influence of life history milestones using age and association patterns on the probability of being a crop raider among wild free ranging male African elephants; we focused on males because female elephants are not known to raid crops in our study population. We examined several features of an elephant association network; network density, community structure and association based on age similarity since they are known to influence the spread of behaviors in a population. We found that older males were more likely to be raiders than younger males, that males were more likely to be raiders when their closest associates were also raiders, and that males were more likely to be raiders when their second closest associates were raiders older than them. The male association network had sparse associations, a tendency for individuals similar in age and raiding status to associate, and a strong community structure. However, raiders were randomly distributed between communities. These features of the elephant association network may limit the spread of raiding behavior and likely determine the prevalence of raiding behavior in elephant populations. Our results suggest that social learning has a major influence on the acquisition of raiding behavior in younger males whereas life history factors are important drivers of raiding behavior in older males. Further, both life-history and network patterns may influence the acquisition and spread of complex behaviors in animal populations and provide insight on managing human-wildlife conflict.     

The crisscrossed agency of a toast: Personhood,individuation and de‐individuation in Luzhou,China

This article addresses debates over individuation in China through consideration of guanxi‐relational feasting in Luzhou, Sichuan. I draw on Ortner's theorisation of subjectivity and agency to probe the often taken‐for‐granted question of cultural personhood which informs social action. Although the social imaginary in Luzhou is increasingly colonised by symbolic individualism, I propose that dominant local notions of personhood and agency, operating within feast practice, continue to define this process. By attending to three aspects of Yan's ‘individualisation thesis’, I demonstrate how local models of person and agency are indispensible to a fuller understanding of social life. Considering the important role ritual speech habits (largely trained in de‐individuating feasting) continue to play in socialising actors to economic institutions and power relationships more generally, individuation in China today remains a largely nominal and aspirational, if symbolically potent and potentially transformative, project.     

Complex Courtship Behavior in the Striped Ground Cricket, Allonemobius socius (Orthoptera: Gryllidae): Does Social Environment Affect Male and Female Behavior?

Complex courtship in the striped ground cricket, Allonemobius socius, involves a series of behaviors alternating between the sexes. We examined if complex courtship allows either or both genders to evaluate their mate and how mating behavior changes in different social environments. While complex courtship may allow discrimination by both sexes, here only females exhibited a preference. Males did not alter their courtship behavior or change spermatophore size for different size females. In contrast, females initiated copulation more quickly with bigger males possessing bigger spermatophores. In a different social environment (additional male, female, or both), males were less likely to omit courtship songs and female discrimination of mates changed. The distinct differences in male and female behavior suggest that subtle changes in social environment can have important consequences in structuring courtship and mating behavior.     

A Bayesian approach to the evolution of social learning

There has been much interest in understanding the evolution of social learning. Investigators have tried to understand when natural selection will favor individuals who imitate others, how imitators should deal with the fact that available models may exhibit different behaviors, and how social and individual learning should interact. In all of this work, social learning and individual learning have been treated as alternative, conceptually distinct processes. Here we present a Bayesian model in which both individual and social learning arise from a single inferential process. Individuals use Bayesian inference to combine social and nonsocial cues about the current state of the environment. This model indicates that natural selection favors individuals who place heavy weight on social cues when the environment changes slowly or when its state cannot be well predicted using nonsocial cues. It also indicates that a conformist bias should be a universal aspect of social learning.