Cultural evolution: implications for understanding the human language faculty and its evolution

Human language is unique among the communication systems of the natural world: it is socially learned and, as a consequence of its recursively compositional structure, offers open-ended communicative potential. The structure of this communication system can be explained as a consequence of the evolution of the human biological capacity for language or the cultural evolution of language itself. We argue, supported by a formal model, that an explanatory account that involves some role for cultural evolution has profound implications for our understanding of the biological evolution of the language faculty: under a number of reasonable scenarios, cultural evolution can shield the language faculty from selection, such that strongly constraining language-specific learning biases are unlikely to evolve. We therefore argue that language is best seen as a consequence of cultural evolution in populations with a weak and/or domain-general language faculty.     

Learning bias, cultural evolution of language, and the biological evolution of the language faculty

The biases of individual language learners act to determine the learnability and cultural stability of languages: learners come to the language learning task with biases which make certain linguistic systems easier to acquire than others. These biases are repeatedly applied during the process of language transmission, and consequently should effect the types of languages we see in human populations. Understanding the cultural evolutionary consequences of particular learning biases is therefore central to understanding the link between language learning in individuals and language universals, common structural properties shared by all the world’s languages. This paper reviews a range of models and experimental studies which show that weak biases in individual learners can have strong effects on the structure of socially learned systems such as language, suggesting that strong universal tendencies in language structure do not require us to postulate strong underlying biases or constraints on language learning. Furthermore, understanding the relationship between learner biases and language design has implications for theories of the evolution of those learning biases: models of gene-culture coevolution suggest that, in situations where a cultural dynamic mediates between properties of individual learners and properties of language in this way, biological evolution is unlikely to lead to the emergence of strong constraints on learning.     

Introduction: integrating genetic and cultural evolutionary approaches to language

The papers in this special issue of Human Biology address recent research in the field of language evolution, both the genetic evolution of the language faculty and the cultural evolution of specific languages. While both of these areas have received increasing interest in recent years, there is also a need to integrate these somewhat separate efforts and explore the relevant gene-culture coevolutionary interactions. Here we summarize the individual contributions, set them in the context of the wider literature, and identify outstanding future research questions. The first set of papers concerns the comparative study of nonhuman communication in primates and birds from both a behavioral and neurobiological perspective, revealing evidence for several common language-related traits in various nonhuman species and providing clues as to the evolutionary origin and function of the human language faculty. The second set of papers discusses the consequences of viewing language as a culturally evolving system in its own right, including claims that this removes the need for strong genetic biases for language acquisition, and that phylogenetic evolutionary methods can be used to reconstruct language histories. We conclude by highlighting outstanding areas for future research, including identifying the precise selection pressures that gave rise to the language faculty in ancestral hominin species, and determining the strength, domain specificity, and origin of the cultural transmission biases that shape languages as they pass along successive generations of language learners.     

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.     

The role of genetic biases in shaping the correlations between languages and genes

It has recently been proposed [Dediu, D., Ladd, D.R., 2007. Linguistic tone is related to the population frequency of the adaptive haplogroups of two brain size genes, ASPM and Microcephalin. Proc. Natl Acad. Sci. USA 104(26), 10944-10949] that genetically coded linguistic biases can influence the trajectory of language change. However, the nature of such biases and the conditions under which they can become manifest have remained vague. The present paper explores computationally two plausible types of linguistic acquisition biases in a population of agents implementing realistic genetic, linguistic and demographic processes. One type of bias represents an innate asymmetric initial state (initial expectation bias) while the other an innate asymmetric facility of acquisition (rate of learning bias). It was found that only the second type of bias produces detectable effects on language through cultural transmission across generations and that such effects are produced even by weak biases present at low frequencies in the population. This suggests that learning preference asymmetries, very small at the individual level and not very frequent at the population level, can bias the trajectory of language change through the process of cultural transmission.     

The Biological Origin of Linguistic Diversity

In contrast with animal communication systems, diversity is characteristic of almost every aspect of human language. Languages variously employ tones, clicks, or manual signs to signal differences in meaning; some languages lack the noun-verb distinction (e.g., Straits Salish), whereas others have a proliferation of fine-grained syntactic categories (e.g., Tzeltal); and some languages do without morphology (e.g., Mandarin), while others pack a whole sentence into a single word (e.g., Cayuga). A challenge for evolutionary biology is to reconcile the diversity of languages with the high degree of biological uniformity of their speakers. Here, we model processes of language change and geographical dispersion and find a consistent pressure for flexible learning, irrespective of the language being spoken. This pressure arises because flexible learners can best cope with the observed high rates of linguistic change associated with divergent cultural evolution following human migration. Thus, rather than genetic adaptations for specific aspects of language, such as recursion, the coevolution of genes and fast-changing linguistic structure provides the biological basis for linguistic diversity. Only biological adaptations for flexible learning combined with cultural evolution can explain how each child has the potential to learn any human language.     

Incidental Acquisition of Foreign Language Vocabulary through Brief Multi-Modal Exposure

First language acquisition requires relatively little effort compared to foreign language acquisition and happens more naturally through informal learning. Informal exposure can also benefit foreign language learning, although evidence for this has been limited to speech perception and production. An important question is whether informal exposure to spoken foreign language also leads to vocabulary learning through the creation of form-meaning links. Here we tested the impact of exposure to foreign language words presented with pictures in an incidental learning phase on subsequent explicit foreign language learning. In the explicit learning phase, we asked adults to learn translation equivalents of foreign language words, some of which had appeared in the incidental learning phase. Results revealed rapid learning of the foreign language words in the incidental learning phase showing that informal exposure to multi-modal foreign language leads to foreign language vocabulary acquisition. The creation of form-meaning links during the incidental learning phase is discussed.     

Cultural selection drives the evolution of human communication systems

Human communication systems evolve culturally, but the evolutionary mechanisms that drive this evolution are not well understood. Against a baseline that communication variants spread in a population following neutral evolutionary dynamics (also known as drift models), we tested the role of two cultural selection models: coordination- and content-biased. We constructed a parametrized mixed probabilistic model of the spread of communicative variants in four 8-person laboratory micro-societies engaged in a simple communication game. We found that selectionist models, working in combination, explain the majority of the empirical data. The best-fitting parameter setting includes an egocentric bias and a content bias, suggesting that participants retained their own previously used communicative variants unless they encountered a superior (content-biased) variant, in which case it was adopted. This novel pattern of results suggests that (i) a theory of the cultural evolution of human communication systems must integrate selectionist models and (ii) human communication systems are functionally adaptive complex systems.     

The evolutionary language game: an orthogonal approach

Evolutionary game dynamics have been proposed as a mathematical framework for the cultural evolution of language and more specifically the evolution of vocabulary. This article discusses a model that is mutually exclusive in its underlying principals with some previously suggested models. The model describes how individuals in a population culturally acquire a vocabulary by actively participating in the acquisition process instead of passively observing and communicate through peer-to-peer interactions instead of vertical parent-offspring relations. Concretely, a notion of social/cultural learning called the naming game is first abstracted using learning theory. This abstraction defines the required cultural transmission mechanism for an evolutionary process. Second, the derived transmission system is expressed in terms of the well-known selection-mutation model defined in the context of evolutionary dynamics. In this way, the analogy between social learning and evolution at the level of meaning-word associations is made explicit. Although only horizontal and oblique transmission structures will be considered, extensions to vertical structures over different genetic generations can easily be incorporated. We provide a number of simplified experiments to clarify our reasoning.     

The evolutionary language game.

We explore how evolutionary game dynamics have to be modified to accomodate a mathematical framework for the evolution of language. In particular, we are interested in the evolution of vocabulary, that is associations between signals and objects. We assume that successful communication contributes to biological fitness: individuals who communicate well leave more offspring. Children inherit from their parents a strategy for language learning (a language acquisition device). We consider three mechanisms whereby language is passed from one generation to the next: (i) parental learning: children learn the language of their parents; (ii) role model learning: children learn the language of individuals with a high payoff; and (iii) random learning: children learn the language of randomly chosen individuals. We show that parental and role model learning outperform random learning. Then we introduce mistakes in language learning and study how this process changes language over time. Mistakes increase the overall efficacy of parental and role model learning: in a world with errors evolutionary adaptation is more efficient. Our model also provides a simple explanation why homonomy is common while synonymy is rare.     

The bridge of iconicity: from a world of experience to the experience of language

Iconicity, a resemblance between properties of linguistic form (both in spoken and signed languages) and meaning, has traditionally been considered to be a marginal, irrelevant phenomenon for our understanding of language processing, development and evolution. Rather, the arbitrary and symbolic nature of language has long been taken as a design feature of the human linguistic system. In this paper, we propose an alternative framework in which iconicity in face-to-face communication (spoken and signed) is a powerful vehicle for bridging between language and human sensori-motor experience, and, as such, iconicity provides a key to understanding language evolution, development and processing. In language evolution, iconicity might have played a key role in establishing displacement (the ability of language to refer beyond what is immediately present), which is core to what language does; in ontogenesis, iconicity might play a critical role in supporting referentiality (learning to map linguistic labels to objects, events, etc., in the world), which is core to vocabulary development. Finally, in language processing, iconicity could provide a mechanism to account for how language comes to be embodied (grounded in our sensory and motor systems), which is core to meaningful communication.     

Review. Theoretical and empirical evidence for the impact of inductive biases on cultural evolution

The question of how much the outcomes of cultural evolution are shaped by the cognitive capacities of human learners has been explored in several disciplines, including psychology, anthropology and linguistics. We address this question through a detailed investigation of transmission chains, in which each person passes information to another along a chain. We review mathematical and empirical evidence that shows that under general conditions, and across experimental paradigms, the information passed along transmission chains will be affected by the inductive biases of the people involved-the constraints on learning and memory, which influence conclusions from limited data. The mathematical analysis considers the case where each person is a rational Bayesian agent. The empirical work consists of behavioural experiments in which human participants are shown to operate in the manner predicted by the Bayesian framework. Specifically, in situations in which each person's response is used to determine the data seen by the next person, people converge on concepts consistent with their inductive biases irrespective of the information seen by the first member of the chain. We then relate the Bayesian analysis of transmission chains to models of biological evolution, clarifying how chains of individuals correspond to population-level models and how selective forces can be incorporated into our models. Taken together, these results indicate how laboratory studies of transmission chains can provide information about the dynamics of cultural evolution and illustrate that inductive biases can have a significant impact on these dynamics.     

Early language acquisition: cracking the speech code

Infants learn language with remarkable speed, but how they do it remains a mystery. New data show that infants use computational strategies to detect the statistical and prosodic patterns in language input, and that this leads to the discovery of phonemes and words. Social interaction with another human being affects speech learning in a way that resembles communicative learning in songbirds. The brain's commitment to the statistical and prosodic patterns that are experienced early in life might help to explain the long-standing puzzle of why infants are better language learners than adults. Successful learning by infants, as well as constraints on that learning, are changing theories of language acquisition.     

Small worlds and semantic network growth in typical and late talkers

Network analysis has demonstrated that systems ranging from social networks to electric power grids often involve a small world structure-with local clustering but global ac cess. Critically, small world structure has also been shown to characterize adult human semantic networks. Moreover, the connectivity pattern of these mature networks is consistent with lexical growth processes in which children add new words to their vocabulary based on the structure of the language-learning environment. However, thus far, there is no direct evidence that a child's individual semantic network structure is associated with their early language learning. Here we show that, while typically developing children's early networks show small world structure as early as 15 months and with as few as 55 words, children with language delay (late talkers) have this structure to a smaller degree. This implicates a maladaptive bias in word acquisition for late talkers, potentially indicating a preference for "oddball" words. The findings provide the first evidence of a link between small-world connectivity and lexical development in individual children.     

Genetic biasing through cultural transmission: Do simple Bayesian models of language evolution generalise?

The recent Bayesian approaches to language evolution and change seem to suggest that genetic biases can impact on the characteristics of language, but, at the same time, that its cultural transmission can partially free it from these same genetic constraints. One of the current debates centres on the striking differences between sampling and a posteriori maximising Bayesian learners, with the first converging on the prior bias while the latter allows a certain freedom to language evolution. The present paper shows that this difference disappears if populations more complex than a single teacher and a single learner are considered, with the resulting behaviours more similar to the sampler. This suggests that generalisations based on the language produced by Bayesian agents in such homogeneous single agent chains are not warranted. It is not clear which of the assumptions in such models are responsible, but these findings seem to support the rising concerns on the validity of the “acquisitionist” assumption, whereby the locus of language change and evolution is taken to be the first language acquirers (children) as opposed to the competent language users (the adults).     

Chaos and language

Human language is a complex communication system with unlimited expressibility. Children spontaneously develop a native language by exposure to linguistic data from their speech community. Over historical time, languages change dramatically and unpredictably by accumulation of small changes and by interaction with other languages. We have previously developed a mathematical model for the acquisition and evolution of language in heterogeneous populations of speakers. This model is based on game dynamical equations with learning. Here, we show that simple examples of such equations can display complex limit cycles and chaos. Hence, language dynamical equations mimic complicated and unpredictable changes of languages over time. In terms of evolutionary game theory, we note that imperfect learning can induce chaotic switching among strict Nash equilibria.     

Manipulations of sensory experiences during development reveal mechanisms underlying vocal learning biases in zebra finches

Biological predispositions in learning can bias and constrain the cultural evolution of social and communicative behaviors (e.g., speech and birdsong), and lead to the emergence of behavioral and cultural “universals.” For example, surveys of laboratory and wild populations of zebra finches (Taeniopygia guttata) document consistent patterning of vocal elements (“syllables”) with respect to their acoustic properties (e.g., duration, mean frequency). Furthermore, such universal patterns are also produced by birds that are experimentally tutored with songs containing randomly sequenced syllables (“tutored birds”). Despite extensive demonstrations of learning biases, much remains to be uncovered about the nature of biological predispositions that bias song learning and production in songbirds. Here, we examined the degree to which “innate” auditory templates and/or biases in vocal motor production contribute to vocal learning biases and production in zebra finches. Such contributions can be revealed by examining acoustic patterns in the songs of birds raised without sensory exposure to song (“untutored birds”) or of birds that are unable to hear from early in development (“early‐deafened birds”). We observed that untutored zebra finches and early‐deafened zebra finches produce songs with positional variation in some acoustic features (e.g., mean frequency) that resemble universal patterns observed in tutored birds. Similar to tutored birds, early‐deafened birds also produced song motifs with alternation in acoustic features across adjacent syllables. That universal acoustic patterns are observed in the songs of both untutored and early‐deafened birds highlights the contribution motor production biases to the emergence of universals in culturally transmitted behaviors.     

Biases in cultural transmission shape the turnover of popular traits

The neutral model of cultural evolution, which assumes that copying is unbiased, provides precise predictions regarding frequency distributions of traits and the turnover within a popularity-ranked list. Here we study turnover in ranked lists and identify where the turnover departs from neutral model predictions to detect transmission biases in three different domains: color terms usage in English language 20th century books, popularity of early (1880–1930) and recent (1960–2010) USA baby names, and musical preferences of users of the Web site Last.fm. To help characterize the type of transmission bias, we modify the neutral model to include a content-based bias and two context-based biases (conformity and anti-conformity). How these modified models match real data helps us to infer, from population scale observations, when cultural transmission is biased, and, to some extent, what kind of biases are operating at individual level.     

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.     

Social learning across the life cycle: cultural knowledge acquisition for honey collection among the Jenu Kuruba,India

Accounting for age-dependent patterns of knowledge transmission is critical for understanding cultural evolution in age-structured populations. Cultural evolution theory predicts which social learning biases we expect people to use, but much less often when—during a person's life cycle— different social learning biases will be used. By measuring knowledge and skill variation among age cohorts, it is possible to infer how people socially acquire different types of knowledge at different ages. We use this strategy among the Jenu Kuruba, a tribal community in South India. We document the accumulation of local knowledge required for collecting wild honey among 71 children and 125 adults from five communities. Combining quantitative measurements of knowledge with measures of four honey-collecting skills and self-reported data on learning age, we infer patterns of social learning across the lifecycle. We find that (1) most knowledge related to honey collecting is acquired by the early 20s, and later social learning mainly functions to update information; (2) the eldest cohort has the highest average explicit knowledge, although the most knowledgeable or skilled individuals do not always belong to the most elderly cohort; (3) length of learning can be affected by age-dependent trade-offs of costs and benefits to learners; and (4) children tend to learn from parents, but individuals use other demonstrators later in life. These results have implications for current models of cultural evolution.