Runaway cultural niche construction

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.     

Individual acquisition of “stick pounding” behavior by naïve chimpanzees

Many studies investigating culture in nonhuman animals tend to focus on the inferred need of social learning mechanisms that transmit the form of a behavior to explain the population differences observed in wild animal behavioral repertoires. This research focus often results in studies overlooking the possibility of individuals being able to develop behavioral forms without requiring social learning. The disregard of individual learning abilities is most clearly observed in the nonhuman great ape literature, where there is a persistent claim that chimpanzee behaviors, in particular, require various forms of social learning mechanisms. These special social learning abilities have been argued to explain the acquisition of the relatively large behavioral repertoires observed across chimpanzee populations. However, current evidence suggests that although low‐fidelity social learning plays a role in harmonizing and stabilizing the frequency of behaviors within chimpanzee populations, some (if not all) of the forms that chimpanzee behaviors take may develop independently of social learning. If so, they would be latent solutions—behavioral forms that can (re‐)emerge even in the absence of observational opportunities, via individual (re)innovations. Through a combination of individual and low‐fidelity social learning, the population‐wide patterns of behaviors observed in great ape species are then established and stably maintained. This is the Zone of Latent Solutions (ZLS) hypothesis. The current study experimentally tested the ZLS hypothesis for pestle pounding, a wild chimpanzee behavior. We tested the reinnovation of this behavior in semi‐wild chimpanzees at Chimfunshi Wildlife Orphanage in Zambia, Africa, (N = 90, tested in four social groups). Crucially, all subjects were naïve to stick pounding before testing. Three out of the four tested groups reinnovated stick pounding—clearly demonstrating that this behavioral form does not require social learning. These findings provide support for the ZLS hypothesis alongside further evidence for the individual learning abilities of chimpanzees.     

Genes,culture, and the human niche: An overview

The sharp distinction between biological traits and culturally based traits, which had long been standard in evolutionary approaches to behavior, was blurred in the early 1980s by mathematical models that allowed a co‐dependent evolution of genetic transmission and cultural information. Niche‐construction theory has since added another contrast to standard evolutionary theory, in that it views niche construction as a cause of evolutionary change rather than simply a product of selection. While offering a new understanding of the coevolution of genes, culture, and human behavior, niche‐construction models also invoke multivariate causality, which require multiple time series to resolve. The empirical challenge lies in obtaining time‐series data on causal pathways involved in the coevolution of genes, culture, and behavior. This is a significant issue in archeology, where time series are often sparse and causal behaviors are represented only by proxies in the material record.     

Traditions in monkeys

Traditions, defined as relatively long‐lasting behavioral practices shared among members of a group partly via social learning, were studied in monkeys, specifically Japanese macaques, before being studied in great apes. Although apes and humans may share some social learning capacities that are absent in monkeys, a complete understanding of the roots of human culture requires attention to the socioecological conditions favoring traditions, however generated, in animals generally and in multiple behavioral domains. Using the four criteria of intergroup variation, observation of the origin and spread of a novel behavior, dissemination patterned according to age or kinship, or individuals' close observation of others' performance of the behavior before engaging in it, over twenty‐five behaviors have been nominated as traditions in free‐ranging monkey populations. Tolerant gregariousness has been proposed to increase the likelihood of the emergence of traditions in any behavioral domain. Omnivory and extractive foraging should favor the emergence of foraging‐related traditions; strong cooperative relationships and flexible coalitionary structure should favor the emergence of social conventions that function to test social bonds. These conditions are taxonomically widespread. Thus, the current restriction of most reported free‐ranging monkey traditions to two taxa (Macaca fuscata and Cebus) is likely to reflect variation in primatologists' research goals, methods, and concepts rather than real interspecific variation in the propensity to generate traditions.     

Rapid evolution of social learning

Culture is widely thought to be beneficial when social learning is less costly than individual learning and thus may explain the enormous ecological success of humans. Rogers (1988. Does biology constrain culture. Am. Anthropol.  90 : 819–831) contradicted this common view by showing that the evolution of social learning does not necessarily increase the net benefits of learned behaviours in a variable environment. Using simulation experiments, we re‐analysed extensions of Rogers’ model after relaxing the assumption that genetic evolution is much slower than cultural evolution. Our results show that this assumption is crucial for Rogers’ finding. For many parameter settings, genetic and cultural evolution occur on the same time scale, and feedback effects between genetic and cultural dynamics increase the net benefits. Thus, by avoiding the costs of individual learning, social learning can increase ecological success. Furthermore, we found that rapid evolution can limit the evolution of complex social learning strategies, which have been proposed to be widespread in animals.     

Conservation and management of the culture of bears

Culture is widely accepted as an important social factor present across a wide range of species. Bears have a culture as defined as behavioral traditions inherited through social learning usually from mothers to offspring. Successful bear cultures can enhance fitness and resource exploitation benefits. In contrast, some bear cultures related to response to humans and human‐related foods can be maladaptive and result in reduced fitness and direct mortality. In environments with minimal human influence most bear culture has evolved over generations to be beneficial and well adapted to enhance fitness. However, most bears across the world do not live in areas with minimal human influence and in these areas, bear culture is often changed by bear interactions with humans, usually to the detriment of bear survival. We highlight the importance of identifying unique bear cultural traits that allow efficient use of local resources and the value of careful management to preserve these adaptive cultural behaviors. It is also important to select against maladaptive cultural behaviors that are usually related to humans in order to reduce human–bear conflicts and high bear mortality. We use examples from Yellowstone National Park to demonstrate how long‐term management to reduce maladaptive bear cultures related to humans has resulted in healthy bear populations and a low level of human–bear conflict in spite of a high number of Yellowstone National Park visitors in close association with bears.     

Conformity in mate choice,the overlooked social component of animal and human culture

Although conformity as a major driver for human cultural evolution is a well-accepted and intensely studied phenomenon, its importance for non-human animal culture has been largely overlooked until recently. This limited for decades the possibility of studying the roots of human culture. Here, we provide a historical review of the study of conformity in both humans and non-human animals. We identify gaps in knowledge and propose an evolutionary route towards the sophisticated cultural processes that characterize humanity. A landmark in the study of conformity is Solomon Asch's famous experiment on humans in 1955. By contrast, interest in conformity among evolutionary biologists has only become salient since the turn of the new millennium. A striking result of our review is that, although studies of conformity have examined many biological contexts, only one looked at mate choice. This is surprising because mate choice is probably the only context in which conformity has self-reinforcing advantages across generations. Within a metapopulation, i.e. a group of subpopulations connected by dispersing individuals, dispersers able to conform to the local preference for a given type of mate have a strong and multigenerational fitness advantage. This is because once females within one subpopulation locally show a bias for one type of males, immigrant females who do not conform to the local trend have sons, grandsons, etc. of the non-preferred phenotype, which negatively and cumulatively affects fitness over generations in a process reminiscent of the Fisher runaway process. This led us to suggest a sex-driven origin of conformity, indicating a possible evolutionary route towards animal and human culture that is rooted in the basic, and thus ancient, social constraints acting on mating preferences within a metapopulation. In a generic model, we show that dispersal among subpopulations within a metapopulation can effectively maintain independent Fisher runaway processes within subpopulations, while favouring the evolution of social learning and conformity at the metapopulation scale; both being essential for the evolution of long-lasting local traditions. The proposed evolutionary route to social learning and conformity casts surprising light on one of the major processes that much later participated in making us human. We further highlight several research avenues to define the spectrum of conformity better, and to account for its complexity. Future studies of conformity should incorporate experimental manipulation of group majority. We also encourage the study of potential links between conformity and mate copying, animal aggregations, and collective actions. Moreover, validation of the sex-driven origin of conformity will rest on the capacity of human and evolutionary sciences to investigate jointly the origin of social learning and conformity. This constitutes a stimulating common agenda and militates for a rapprochement between these two currently largely independent research areas.     

The question of animal culture

In this paper I consider whether traditional behaviors of animals, like traditions of humans, are transmitted by imitation learning. Review of the literature on problem solving by captive primates, and detailed consideration of two widely cited instances of purported learning by imitation and of culture in free-living primates (sweet-potato washing by Japanese macaques and termite fishing by chimpanzees), suggests that nonhuman primates do not learn to solve problems by imitation. It may, therefore, be misleading to treat animal traditions and human culture as homologous (rather than analogous) and to refer to animal traditions as cultural.     

Human cumulative culture: a comparative perspective

Many animals exhibit social learning and behavioural traditions, but human culture exhibits unparalleled complexity and diversity, and is unambiguously cumulative in character. These similarities and differences have spawned a debate over whether animal traditions and human culture are reliant on homologous or analogous psychological processes. Human cumulative culture combines high‐fidelity transmission of cultural knowledge with beneficial modifications to generate a ‘ratcheting’ in technological complexity, leading to the development of traits far more complex than one individual could invent alone. Claims have been made for cumulative culture in several species of animals, including chimpanzees, orangutans and New Caledonian crows, but these remain contentious. Whilst initial work on the topic of cumulative culture was largely theoretical, employing mathematical methods developed by population biologists, in recent years researchers from a wide range of disciplines, including psychology, biology, economics, biological anthropology, linguistics and archaeology, have turned their attention to the experimental investigation of cumulative culture. We review this literature, highlighting advances made in understanding the underlying processes of cumulative culture and emphasising areas of agreement and disagreement amongst investigators in separate fields.     

Vocal culture in New Caledonian crows Corvus moneduloides

While recent research suggests that some animal species may possess ‘cultural’ traditions, much of the current evidence for wild populations remains contentious. This is largely due to the difficulty of demonstrating a fundamental prerequisite for the existence of culture: social learning. As the only case where social learning has been demonstrated conclusively, and subsequently linked to spatial or temporal trait variation, avian vocal dialects are the best studied, and most widely accepted, form of animal culture. Here, we investigate the potential for vocal culture in one of the few animals for which material culture has been suggested: the New Caledonian crow Corvus moneduloides. We show that this species: (1) possesses the capacity for social learning of vocalizations (experimental evidence in the form of a captive subject that reproduces human speech and other anthropogenic noises); and (2) exhibits significant large‐scale, population‐level variation in its vocalizations (cross‐island playback experiments, with analyses controlling for a substantial set of potentially confounding variables). In combination, this provides strong evidence for the existence of ‘culture’ in these birds. More specifically, our findings reveal that the species exhibits sufficient social learning mechanisms, and within‐population structuring, to generate and perpetuate cultural variation in at least one behavioural domain. This is a critical first step towards demonstrating cultural transmission in other behaviours, including tool manufacture and tool use, opening the door for the simultaneous investigation of vocal and material culture in a nonhuman species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 767–776.     

Origins of altruism diversity ii: runaway coevolution of altruistic strategies via "reciprocal niche construction"

Understanding the evolution of altruism requires knowledge of both its constraints and its drivers. Here we show that, paradoxically, ecological constraints on altruism may ultimately be its strongest driver. We construct a two-trait, coevolutionary adaptive dynamics model of social evolution in a genetically structured population with local resource competition. The intensity of local resource competition, which influences the direction and strength of social selection and which is typically treated as a static parameter, is here allowed to be an evolvable trait. Evolution of survival/fecundity altruism, which requires weak local competition, increases local competition as it evolves, creating negative environmental feedback that ultimately inhibits its further evolutionary advance. Alternatively, evolution of resource-based altruism, which requires strong local competition, weakens local competition as it evolves, also ultimately causing its own evolution to stall. When evolving independently, these altruistic strategies are intrinsically self-limiting. However, the coexistence of these two altruism types transforms the negative ecoevolutionary feedback generated by each strategy on itself into positive feedback on the other, allowing the presence of one trait to drive the evolution of the other. We call this feedback conversion "reciprocal niche construction." In the absence of constraints, this process leads to runaway coevolution of altruism types. We discuss applications to the origins and evolution of eusociality, division of labor, the inordinate ecological success of eusocial species, and the interaction between technology and demography in human evolution. Our theory suggests that the evolution of extreme sociality may often be an autocatalytic process.     

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.     

Runaway sexual selection when female preferences are directly selected

We introduce models for the runaway coevolution of female mating preferences and male display traits. The models generalize earlier results by allowing for direct natural selection on the preference, arbitrary forms of mate choice, and fairly general assumptions about the underlying genetics. Results show that a runaway is less likely when there is direct selection on the preference, but that it is still possible if there is a sufficiently large phenotypic correlation between the female's preference and the male's trait among mated pairs. Comparison of three preference functions introduced by Lande (1981) shows that open-ended preferences are particularly prone to a runaway, and that absolute preferences require very large differences between females in their preferences. We analyze the causes of the runaway seen in a model developed by Iwasa and Pomiankowski (1995).     

The biology of cultural conflict

Although culture is usually thought of as the collection of knowledge and traditions that are transmitted outside of biology, evidence continues to accumulate showing how biology and culture are inseparably intertwined. Cultural conflict will occur only when the beliefs and traditions of one cultural group represent a challenge to individuals of another. Such a challenge will elicit brain processes involved in cognitive decision-making, emotional activation and physiological arousal associated with the outbreak, conduct and resolution of conflict. Key targets to understand bio-cultural differences include primitive drives-how the brain responds to likes and dislikes, how it discounts the future, and how this relates to reproductive behaviour-but also higher level functions, such as how the mind represents and values the surrounding physical and social environment. Future cultural wars, while they may bear familiar labels of religion and politics, will ultimately be fought over control of our biology and our environment.     

Social enhancement can create adaptive,arbitrary and maladaptive cultural traditions

Many animals are known to learn socially, i.e. they are able to acquire new behaviours by using information from other individuals. Researchers distinguish between a number of different social-learning mechanisms such as imitation and social enhancement. Social enhancement is a simple form of social learning that is among the most widespread in animals. However, unlike imitation, it is debated whether social enhancement can create cultural traditions. Based on a recent study on capuchin monkeys, we developed an agent-based model to test the hypotheses that (i) social enhancement can create and maintain stable traditions and (ii) social enhancement can create cultural conformity. Our results supported both hypotheses. A key factor that led to the creation of cultural conformity and traditions was the repeated interaction of individual reinforcement and social enhancement learning. This result emphasizes that the emergence of cultural conformity does not necessarily require cognitively complex mechanisms such as ‘copying the majority’ or group norms. In addition, we observed that social enhancement can create learning dynamics similar to a ‘copy when uncertain’ learning strategy. Results from additional analyses also point to situations that should favour the evolution of learning mechanisms more sophisticated than social enhancement.     

Exploring gene-culture interactions: insights from handedness, sexual selection and niche-construction case studies

Genes and culture represent two streams of inheritance that for millions of years have flowed down the generations and interacted. Genetic propensities, expressed throughout development, influence what cultural organisms learn. Culturally transmitted information, expressed in behaviour and artefacts, spreads through populations, modifying selection acting back on populations. Drawing on three case studies, I will illustrate how this gene-culture coevolution has played a critical role in human evolution. These studies explore (i) the evolution of handedness, (ii) sexual selection with a culturally transmitted mating preference, and (iii) cultural niche construction and human evolution. These analyses shed light on how genes and culture shape each other, and on the significance of feedback mechanisms between biological and cultural processes.     

THE EVOLUTION OF PHENOTYPIC CORRELATIONS AND “DEVELOPMENTAL MEMORY”

Development introduces structured correlations among traits that may constrain or bias the distribution of phenotypes produced. Moreover, when suitable heritable variation exists, natural selection may alter such constraints and correlations, affecting the phenotypic variation available to subsequent selection. However, exactly how the distribution of phenotypes produced by complex developmental systems can be shaped by past selective environments is poorly understood. Here we investigate the evolution of a network of recurrent nonlinear ontogenetic interactions, such as a gene regulation network, in various selective scenarios. We find that evolved networks of this type can exhibit several phenomena that are familiar in cognitive learning systems. These include formation of a distributed associative memory that can “store” and “recall” multiple phenotypes that have been selected in the past, recreate complete adult phenotypic patterns accurately from partial or corrupted embryonic phenotypes, and “generalize” (by exploiting evolved developmental modules) to produce new combinations of phenotypic features. We show that these surprising behaviors follow from an equivalence between the action of natural selection on phenotypic correlations and associative learning, well‐understood in the context of neural networks. This helps to explain how development facilitates the evolution of high‐fitness phenotypes and how this ability changes over evolutionary time.     

Evolution of cultural communication systems: the coevolution of cultural signals and genes encoding learning preferences

In several communication systems that rely on social learning, such as bird song, and possibly human language, the range of signals that can be learned is limited by perceptual biases--predispositions--that are presumably based on genes. In this paper, we examine the coevolution of such genes with the culturally transmitted communication traits themselves, using deterministic population genetic models. We argue that examining how restrictive genetic predispositions are is a useful way of examining the evolutionary origin and maintenance of learning. Under neutral cultural evolution, where no cultural trait has any inherent advantage over another, there is selection in favour of less restrictive genes (genes that allow a wider range of signals to recognized). In contrast, cultural conformity (where the most common cultural trait is favoured) leads to selection in favour of more restrictive genes.     

Reconciling community ecology with evidence of animal culture: Socially-adapted,localized community dynamics?

A growing body of empirical research suggests many animal species are capable of social learning and even have cultural behavioral traditions. Social learning has implications for community ecology; changes in behavior can lead to changes in inter- and intra-specific (between and within species) interactions. The paper explores possible implications of social learning for ecological community dynamics. Four arguments are made: (1) social learning can result in locally-specific ecological relationships; (2) socially-mediated, locally-specific ecological relationships can have localized indirect interspecific population effects; (3) the involvement of multiple co-existing species in socially learned, locally-specific behavior has the potential to create community-wide effects, including varying levels of stability and instability; and (4) social learning can create new intra- and inter-specific selection pressures on local taxa, potentially resulting in rapid evolution. Implications of all four arguments are discussed in relation to community ecology research and modeling.     

文明的生态学分析与绿色文化

绿色文化即人类与环境共进,实现可持续发展的文化,包括持续农业、持续林业和一切不以牺牲环境为代价的绿色产业.根据西双版纳野外文化生态学调查结果,从绿色文化的高度分析和研究了少数民族的生产、生活习惯和传统宗教文化,为可持续发展和自然资源保护寻求一种民族的根源和传统.