Cultural transmission and the evolution of human behaviour: a general approach based on the Price equation

Transmitted culture can be viewed as an inheritance system somewhat independent of genes that is subject to processes of descent with modification in its own right. Although many authors have conceptualized cultural change as a Darwinian process, there is no generally agreed formal framework for defining key concepts such as natural selection, fitness, relatedness and altruism for the cultural case. Here, we present and explore such a framework using the Price equation. Assuming an isolated, independently measurable culturally transmitted trait, we show that cultural natural selection maximizes cultural fitness, a distinct quantity from genetic fitness, and also that cultural relatedness and cultural altruism are not reducible to or necessarily related to their genetic counterparts. We show that antagonistic coevolution will occur between genes and culture whenever cultural fitness is not perfectly aligned with genetic fitness, as genetic selection will shape psychological mechanisms to avoid susceptibility to cultural traits that bear a genetic fitness cost. We discuss the difficulties with conceptualizing cultural change using the framework of evolutionary theory, the degree to which cultural evolution is autonomous from genetic evolution, and the extent to which cultural change should be seen as a Darwinian process. We argue that the nonselection components of evolutionary change are much more important for culture than for genes, and that this and other important differences from the genetic case mean that different approaches and emphases are needed for cultural than genetic processes.     

Not in their genes: Phenotypic flexibility, behavioural traditions and cultural evolution in wild bonnet macaques

Phenotypic flexibility, or the within-genotype, context-dependent, variation in behaviour expressed by single reproductively mature individuals during their lifetimes, often impart a selective advantage to organisms and profoundly influence their survival and reproduction. Another phenomenon apparently not under direct genetic control is behavioural inheritance whereby higher animals are able to acquire information from the behaviour of others by social learning, and, through their own modified behaviour, transmit such information between individuals and across generations. Behavioural information transfer of this nature thus represents another form of inheritance that operates in many animals in tandem with the more basic genetic system. This paper examines the impact that phenotypic flexibility, behavioural inheritance and socially transmitted cultural traditions may have in shaping the structure and dynamics of a primate society — that of the bonnet macaque (Macaca radiata), a primate species endemic to peninsular India. Three principal issues are considered: the role of phenotypic flexibility in shaping social behaviour, the occurrence of individual behavioural traits leading to the establishment of social traditions, and the appearance of cultural evolution amidst such social traditions. Although more prolonged observations are required, these initial findings suggest that phenotypic plasticity, behavioural inheritance and cultural traditions may be much more widespread among primates than have previously been assumed but may have escaped attention due to a preoccupation with genetic inheritance in zoological thinking. This paper is dedicated with admiration and gratitude to the late Dr Raja Ramanna, a man who held all primates in great esteem.     

Untangling cultural inheritance: language diversity and long-house architecture on the Pacific northwest coast

Many recent studies of cultural inheritance have focused on small-scale craft traditions practised by single individuals, which do not require coordinated participation by larger social collectives. In this paper, we address this gap in the cultural transmission literature by investigating diversity in the vernacular architecture of the Pacific northwest coast, where communities of hunter-fisher-gatherers constructed immense wooden long-houses at their main winter villages. Quantitative analyses of long-house styles along the coastline draw on a range of models and methods from the biological sciences and are employed to test hypotheses relating to basic patterns of macro-scale cultural diversification, and the degree to which the transmission of housing traits has been constrained by the region's numerous linguistic boundaries. The results indicate relatively strong branching patterns of cultural inheritance and also close associations between regional language history and housing styles, pointing to the potentially crucial role played by language boundaries in structuring large-scale patterns of cultural diversification, especially in relation to 'collective' cultural traditions like housing that require substantial inputs of coordinated labour.     

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.     

Inferring population histories using cultural data

The question as to whether cultures evolve in a manner analogous to that of genetic evolution can be addressed by attempting to reconstruct population histories using cultural data. As others have argued, this can only succeed if cultures are isolated enough to maintain and pass on a central core of traditions that can be modified over time. In this study we used a set of cultural data (canoe design traits from Polynesia) to look for the kinds of patterns and relationships normally found in population genetic studies. After developing new techniques to accommodate the peculiarities of cultural data, we were able to infer an ancestral region (Fiji) and a sequence of cultural origins for these Polynesian societies. In addition, we found evidence of cultural exchange, migration and a serial founder effect. Results were stronger when analyses were based on functional traits (presumably subject to natural selection and convergence) rather than symbolic or stylistic traits (probably subject to cultural selection for rapid divergence). These patterns strongly suggest that cultural evolution, while clearly affected by cultural exchange, is also subject to some of the same processes and constraints as genetic evolution.     

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.     

Cumulative cultural dynamics and the coevolution of cultural innovation and transmission: an ESS model for panmictic and structured populations

When individuals in a population can acquire traits through learning, each individual may express a certain number of distinct cultural traits. These traits may have been either invented by the individual himself or acquired from others in the population. Here, we develop a game theoretic model for the accumulation of cultural traits through individual and social learning. We explore how the rates of innovation, decay, and transmission of cultural traits affect the evolutionary stable (ES) levels of individual and social learning and the number of cultural traits expressed by an individual when cultural dynamics are at a steady‐state. We explore the evolution of these phenotypes in both panmictic and structured population settings. Our results suggest that in panmictic populations, the ES level of learning and number of traits tend to be independent of the social transmission rate of cultural traits and is mainly affected by the innovation and decay rates. By contrast, in structured populations, where interactions occur between relatives, the ES level of learning and the number of traits per individual can be increased (relative to the panmictic case) and may then markedly depend on the transmission rate of cultural traits. This suggests that kin selection may be one additional solution to Rogers's paradox of nonadaptive culture.     

Modelling the evolution and diversity of cumulative culture

Previous work on mathematical models of cultural evolution has mainly focused on the diffusion of simple cultural elements. However, a characteristic feature of human cultural evolution is the seemingly limitless appearance of new and increasingly complex cultural elements. Here, we develop a general modelling framework to study such cumulative processes, in which we assume that the appearance and disappearance of cultural elements are stochastic events that depend on the current state of culture. Five scenarios are explored: evolution of independent cultural elements, stepwise modification of elements, differentiation or combination of elements and systems of cultural elements. As one application of our framework, we study the evolution of cultural diversity (in time as well as between groups).     

Studying phenotypic plasticity: the advantages of a broad approach

Human evolution has clearly been shaped by gene–culture interactions, and there is growing evidence that similar processes also act on populations of non‐human animals. Recent theoretical studies have shown that culture can be an important evolutionary mechanism because of the ability of cultural traits to spread rapidly both vertically, obliquely, and horizontally, resulting in decreased within‐group variance and increased between‐group variance. Here, we collate the extensive literature on population divergence in killer whales (Orcinus orca), and argue that they are undergoing ecological speciation as a result of dietary specializations. Although we cannot exclude the possibility that cultural divergence pre‐dates ecological divergence, we propose that cultural differences in the form of learned behaviours between ecologically divergent killer whale populations have resulted in sufficient reproductive isolation even in sympatry to lead to incipient speciation. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 1–17.     

Determinants of echolocation call frequency variation in the Formosan lesser horseshoe bat (Rhinolophus monoceros)

The origin and maintenance of intraspecific variation in vocal signals is important for population divergence and speciation. Where vocalizations are transmitted by vertical cultural inheritance, similarity will reflect co-ancestry, and thus vocal divergence should reflect genetic structure. Horseshoe bats are characterized by echolocation calls dominated by a constant frequency component that is partly determined by maternal imprinting. Although previous studies showed that constant frequency calls are also influenced by some non-genetic factors, it is not known how frequency relates to genetic structure. To test this, we related constant frequency variation to genetic and non-genetic variables in the Formosan lesser horseshoe bat (Rhinolophus monoceros). Recordings of bats from across Taiwan revealed that females called at higher frequencies than males; however, we found no effect of environmental or morphological factors on call frequency. By comparison, variation showed clear population structure, with frequencies lower in the centre and east, and higher in the north and south. Within these regions, frequency divergence was directional and correlated with geographical distance, suggesting that call frequencies are subject to cultural drift. However, microsatellite clustering analysis showed that broad differences in constant frequency among populations corresponded to discontinuities in allele frequencies resulting from vicariant events. Our results provide evidence that the processes shaping genetic subdivision have concomitant consequences for divergence in echolocation call frequency.     

Genetic consequences of breaking migratory traditions in barnacle geese Branta leucopsis

Cultural transmission of migratory traditions enables species to deal with their environment based on experiences from earlier generations. Also, it allows a more adequate and rapid response to rapidly changing environments. When individuals break with their migratory traditions, new population structures can emerge that may affect gene flow. Recently, the migratory traditions of the Barnacle Goose Branta leucopsis changed, and new populations differing in migratory distance emerged. Here, we investigate the population genetic structure of the Barnacle Goose to evaluate the consequences of altered migratory traditions. We used a set of 358 single nucleotide polymorphism (SNP) markers to genotype 418 individuals from breeding populations in Greenland, Spitsbergen, Russia, Sweden and the Netherlands, the latter two being newly emerged populations. We used discriminant analysis of principal components, F_ST, linkage disequilibrium and a comparison of geneflow models using migrate ‐n to show that there is significant population structure, but that relatively many pairs of SNPs are in linkage disequilibrium, suggesting recent admixture between these populations. Despite the assumed traditions of migration within populations, we also show that genetic exchange occurs between all populations. The newly established nonmigratory population in the Netherlands is characterized by high emigration into other populations, which suggests more exploratory behaviour, possibly as a result of shortened parental care. These results suggest that migratory traditions in populations are subject to change in geese and that such changes have population genetic consequences. We argue that the emergence of nonmigration probably resulted from developmental plasticity.     

Transmission fidelity is the key to the build-up of cumulative culture

Many animals have socially transmitted behavioural traditions, but human culture appears unique in that it is cumulative, i.e. human cultural traits increase in diversity and complexity over time. It is often suggested that high-fidelity cultural transmission is necessary for cumulative culture to occur through refinement, a process known as 'ratcheting', but this hypothesis has never been formally evaluated. We discuss processes of information transmission and loss of traits from a cognitive viewpoint alongside other cultural processes of novel invention (generation of entirely new traits), modification (refinement of existing traits) and combination (bringing together two established traits to generate a new trait). We develop a simple cultural transmission model that does not assume major evolutionary changes (e.g. in brain architecture) and show that small changes in the fidelity with which information is passed between individuals can lead to cumulative culture. In comparison, modification and combination have a lesser influence on, and novel invention appears unimportant to, the ratcheting process. Our findings support the idea that high-fidelity transmission is the key driver of human cumulative culture, and that progress in cumulative culture depends more on trait combination than novel invention or trait modification.     

The scope of culture in chimpanzees, humans and ancestral apes

More studies have focused on aspects of chimpanzee behaviour and cognition relevant to the evolution of culture than on any other species except our own. Accordingly, analysis of the features shared by chimpanzees and humans is here used to infer the scope of cultural phenomena in our last common ancestor, at the same time clarifying the nature of the special characteristics that advanced further in the hominin line. To do this, culture is broken down into three major aspects: the large scale, population-level patterning of traditions; social learning mechanisms; and the behavioural and cognitive contents of culture. Each of these is further dissected into subcomponents. Shared features, as well as differences, are identified in as many as a dozen of these, offering a case study for the comparative analysis of culture across animal taxa and a deeper understanding of the roots of our own cultural capacities.     

西南地区文化多样性时空格局

我国西南地区少数民族类别众多,各民族在历史发展中形成了许多优秀的文化传统和独特的生态意识。本研究以县级行政区为基本研究单元,基于民族类别及其人口定量测度了西南地区的文化多样性指数,并比较了1990年、2000年与2010年的动态变化。西南地区文化多样性指数具有明显的空间异质性,整体上呈现西北低东南高的格局。虽然文化多样性指数在时间序列上有一定程度的增加,但总体上县域尺度的文化多样性水平偏低。此外,随着空间尺度的扩大,文化多样性指数逐渐增加,即西南地区整体上表现为"大杂居,小聚居"的分布格局。省域尺度上,文化多样性的差异总和与各组分差异均随时间的推移而减少,同时,其民族构成的相似性趋于增加,表明西南地区各民族人口的分布更加融合,民族交融成为趋势。在西南地区的生态环境背景下,文化多样性的空间分布与平均坡度密切相关。研究探讨了民族融合的社会历史背景,分析了阶梯过渡带具有较高文化多样性的原因,并呼吁加强对西南地区优秀传统文化的重视与保护。     

Cultural Evolution: A Review of Theory,Findings and Controversies

The last two decades have seen an explosion in research analysing cultural change as a Darwinian evolutionary process. Here I provide an overview of the theory of cultural evolution, including its intellectual history, major theoretical tenets and methods, key findings, and prominent criticisms and controversies. ‘Culture’ is defined as socially transmitted information. Cultural evolution is the theory that this socially transmitted information evolves in the manner laid out by Darwin in The Origin of Species, i.e. it comprises a system of variation, differential fitness and inheritance. Cultural evolution is not, however, neo-Darwinian, in that many of the details of genetic evolution may not apply, such as particulate inheritance and random mutation. Following a brief history of this idea, I review theoretical and empirical studies of cultural microevolution, which entails both selection-like processes wherein some cultural variants are more likely to be acquired and transmitted than others, plus transformative processes that alter cultural information during transmission. I also review how phylogenetic methods have been used to reconstruct cultural macroevolution, including the evolution of languages, technology and social organisation. Finally, I discuss recent controversies and debates, including the extent to which culture is proximate or ultimate, the relative role of selective and transformative processes in cultural evolution, the basis of cumulative cultural evolution, the evolution of large-scale human cooperation, and whether social learning is learned or innate. I conclude by highlighting the value of using evolutionary methods to study culture for both the social and biological sciences.     

Culture and courtship in vertebrates: a review of social learning and transmission of courtship systems and mating patterns

Female and male animals often choose mates based upon the complementarity of their courtship behaviours and preferences. The importance of this fact on the evolutionary dynamics of populations has long been appreciated. What has not been appreciated is the role that social learning might play in the transmission of systems of courtship behaviour across generations. This paper addresses the social transmission of courtship behavioural traditions in vertebrates. It discusses views of culture in the context of behavioural signals and preferences in courtship. It then reviews empirical evidence for culture-like processes affecting courtship behaviour, focusing on studies of song learning in passerine birds and work on social learning of mating preferences. The paper concludes with potential future directions for research on social traditions in systems of courtship behaviour, including determining mechanisms of transmission, genetic and non-social environmental effects, and selective factors influencing the stability of behavioural traditions over time. By integrating proximate and ultimate questions for the transmission of courtship systems, this work would increase our understanding of the ways individual development, cultural processes, and population evolution influence, and are in turn influenced by, one another.     

景观传统的现代转译——以北京市中华文化智慧公园为例

在高速城市化和全球化的背景下，传统园林的传承陷入了现实的困境，如何在满足现代城市功能的基础上，继承和发扬传统文化成为研究的热点话题。中华文化智慧公园致力于对景观传统进行转译，并运用数字设计和建造手法，在景观中实现传统文化和现代化的平衡，从而营造出既能符合国际发展趋势，又能体现传统文化内涵的景观。     

The Italian Way to Stem Cell Research: Rethinking the Role of Catholic Religion in Shaping Italian Stem Cell Research Regulations

Stem cell research regulations are highly variable across nations, notwithstanding shared and common ethical concerns. Dominant in political debates has been the so‐called embryo question. However, the permissibility of human embryonic stem cell (hESC) research varies among national regulatory frameworks. Scholars have explained differences by resorting to notions of political culture, traditions of ethical reasoning, discursive strategies and political manoeuvring of involved actors. Explanations based on the role of religion or other cultural structural variables are also employed. This paper analyses the emerging of the Italian regulatory framework on stem cell research using an analytical framework that considers the interplay between cultural structural features, political culture, traditions of ethical reasoning, institutional settings and the discursive and political agency of the actors involved. It aims also to explain the role of Roman Catholic Church in shaping the Italian stem cell research regulation not by treating religion as an autonomous causal factor, but through the analysis of the agency of Catholic and allied actors in the Italian political culture and institutional setting.     

Interactions of genetic and cultural evolution: Models and examples

This paper proposes models and examples of five principal modes of interaction between genes and culture in human evolution. Because genes and culture ultimately interact in the minds of individuals, the models are focused on individual level processes of constrained microevolution. The central hypotheses are (1) that cultural evolution as well as genetic evolution commonly proceeds by the differential transmission of alternative instructions among individuals, (2) that genetic and cultural processes directly interact through mutual influence on each other's differentials of transmission in a population, (3) that the cultural process is often self-selecting by its own criteria, and (4) that these criteria generally operate to enhance rather than oppose human adaptation. Evolutionary change at higher levels, which is particularly important in sociocultural evolution, is interpreted as restructuring the nature and extent of the variability available at the individual level. To clarify the conceptual differences of the models and hopefully to stimulate related analyses in other areas, I discuss selected examples of each of these interactions. I conclude with some remarks on the relative importance of the models to human ecology and evolution.