Dating the origin of language using phonemic diversity

Language is a key adaptation of our species, yet we do not know when it evolved. Here, we use data on language phonemic diversity to estimate a minimum date for the origin of language. We take advantage of the fact that phonemic diversity evolves slowly and use it as a clock to calculate how long the oldest African languages would have to have been around in order to accumulate the number of phonemes they possess today. We use a natural experiment, the colonization of Southeast Asia and Andaman Islands, to estimate the rate at which phonemic diversity increases through time. Using this rate, we estimate that present-day languages date back to the Middle Stone Age in Africa. Our analysis is consistent with the archaeological evidence suggesting that complex human behavior evolved during the Middle Stone Age in Africa, and does not support the view that language is a recent adaptation that has sparked the dispersal of humans out of Africa. While some of our assumptions require testing and our results rely at present on a single case-study, our analysis constitutes the first estimate of when language evolved that is directly based on linguistic data.     

The Role of Phoneme in Mandarin Chinese Production: Evidence from ERPs

Established linguistic theoretical frameworks propose that alphabetic language speakers use phonemes as phonological encoding units during speech production whereas Mandarin Chinese speakers use syllables. This framework was challenged by recent neural evidence of facilitation induced by overlapping initial phonemes, raising the possibility that phonemes also contribute to the phonological encoding process in Chinese. However, there is no evidence of non-initial phoneme involvement in Chinese phonological encoding among representative Chinese speakers, rendering the functional role of phonemes in spoken Chinese controversial. Here, we addressed this issue by systematically investigating the word-initial and non-initial phoneme repetition effect on the electrophysiological signal using a picture-naming priming task in which native Chinese speakers produced disyllabic word pairs. We found that overlapping phonemes in both the initial and non-initial position evoked more positive ERPs in the 180- to 300-ms interval, indicating position-invariant repetition facilitation effect during phonological encoding. Our findings thus revealed the fundamental role of phonemes as independent phonological encoding units in Mandarin Chinese.     

Bringing together linguistic and genetic evidence to test the Bantu expansion

The expansion of Bantu languages represents one of the most momentous events in the history of Africa. While it is well accepted that Bantu languages spread from their homeland (Cameroon/Nigeria) approximately 5000 years ago (ya), there is no consensus about the timing and geographical routes underlying this expansion. Two main models of Bantu expansion have been suggested: The 'early-split' model claims that the most recent ancestor of Eastern languages expanded north of the rainforest towards the Great Lakes region approximately 4000 ya, while the 'late-split' model proposes that Eastern languages diversified from Western languages south of the rainforest approximately 2000 ya. Furthermore, it is unclear whether the language dispersal was coupled with the movement of people, raising the question of language shift versus demic diffusion. We use a novel approach taking into account both the spatial and temporal predictions of the two models and formally test these predictions with linguistic and genetic data. Our results show evidence for a demic diffusion in the genetic data, which is confirmed by the correlations between genetic and linguistic distances. While there is little support for the early-split model, the late-split model shows a relatively good fit to the data. Our analyses demonstrate that subsequent contact among languages/populations strongly affected the signal of the initial migration via isolation by distance.     

Evidence for Direct Geographic Influences on Linguistic Sounds: The Case of Ejectives

We present evidence that the geographic context in which a language is spoken may directly impact its phonological form. We examined the geographic coordinates and elevations of 567 language locations represented in a worldwide phonetic database. Languages with phonemic ejective consonants were found to occur closer to inhabitable regions of high elevation, when contrasted to languages without this class of sounds. In addition, the mean and median elevations of the locations of languages with ejectives were found to be comparatively high. The patterns uncovered surface on all major world landmasses, and are not the result of the influence of particular language families. They reflect a significant and positive worldwide correlation between elevation and the likelihood that a language employs ejective phonemes. In addition to documenting this correlation in detail, we offer two plausible motivations for its existence. We suggest that ejective sounds might be facilitated at higher elevations due to the associated decrease in ambient air pressure, which reduces the physiological effort required for the compression of air in the pharyngeal cavity–a unique articulatory component of ejective sounds. In addition, we hypothesize that ejective sounds may help to mitigate rates of water vapor loss through exhaled air. These explications demonstrate how a reduction of ambient air density could promote the usage of ejective phonemes in a given language. Our results reveal the direct influence of a geographic factor on the basic sound inventories of human languages.     

Bayesian phylogenetic analysis supports an agricultural origin of Japonic languages

Languages, like genes, evolve by a process of descent with modification. This striking similarity between biological and linguistic evolution allows us to apply phylogenetic methods to explore how languages, as well as the people who speak them, are related to one another through evolutionary history. Language phylogenies constructed with lexical data have so far revealed population expansions of Austronesian, Indo-European and Bantu speakers. However, how robustly a phylogenetic approach can chart the history of language evolution and what language phylogenies reveal about human prehistory must be investigated more thoroughly on a global scale. Here we report a phylogeny of 59 Japonic languages and dialects. We used this phylogeny to estimate time depth of its root and compared it with the time suggested by an agricultural expansion scenario for Japanese origin. In agreement with the scenario, our results indicate that Japonic languages descended from a common ancestor approximately 2182 years ago. Together with archaeological and biological evidence, our results suggest that the first farmers of Japan had a profound impact on the origins of both people and languages. On a broader level, our results are consistent with a theory that agricultural expansion is the principal factor for shaping global linguistic diversity.     

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.     

The measurement of linguistic change

Summary Two methods for measuring the amount of linguistic change are described: 1. It is possible to measure the correspondence between two phoneme inventories of two periods in a language's development in the same way as in information theory between the inventories of the transmitted and received symbols (measure of correspondence B). 2. This first method, however, does not measure the development of a phonologic pattern. Therefore the correlation of similarity is computed (as applicated in the psychology of development).     

Patterns of covert speech behavior and phonetic coding

In previous research a discriminative relationship has been established between patterns of covert speech behavior and the phonemic system when processing continuous linguistic material. The goal of the present research was to be more analytic and pinpoint covert neuromuscular speech patterns when one processes specific instances of phonemes. Electromyographic (EMG) recording indicated that the lips are significantly active when visually processing the letter "P"(an instance of bilabial material), but not when processing the letter "T" or a nonlinguistic control (C) stimulus. Similarly, the tongue is significantly active when processing the letter "T" (an instance of lingual-alveolar material), but not when processing the letters "P" or "C". It is concluded that the speech musculature covertly responds systematically as a function of class of phoneme being processed. These results accord with our model that semantic processing ("understanding") occurs when the speech (and other) musculature interacts with linguistic regions of the brain. In the interactions phonetic coding is generated and transmitted through neuromuscular circuits that have cybernetic characteristics.     

Oceanic barriers promote language diversification in the Japanese Islands

Good barriers make good languages. Scholars have long speculated that geographical barriers impede linguistic contact between speech communities and promote language diversification in a manner similar to the process of allopatric speciation. This hypothesis, however, has seldom been tested systematically and quantitatively. Here, we adopt methods from evolutionary biology and attempt to quantify the influence of oceanic barriers on the degree of lexical diversity in the Japanese Islands. Measuring the degree of beta diversity from basic vocabularies, we find that geographical proximity and, more importantly, isolation by surrounding ocean, independently explains a significant proportion of lexical variation across Japonic languages. Further analyses indicate that our results are neither a by‐product of using a distance matrix derived from a Bayesian language phylogeny nor an epiphenomenon of accelerated evolutionary rates in languages spoken by small communities. Moreover, we find that the effect of oceanic barriers is reproducible with the Ainu languages, indicating that our analytic approach as well as the results can be generalized beyond Japonic language family. The findings we report here are the first quantitative evidence that physical barriers formed by ocean can influence language diversification and points to an intriguing common mechanism between linguistic and biological evolution.     

Genetic and linguistic coevolution in Northern Island Melanesia

Recent studies have detailed a remarkable degree of genetic and linguistic diversity in Northern Island Melanesia. Here we utilize that diversity to examine two models of genetic and linguistic coevolution. The first model predicts that genetic and linguistic correspondences formed following population splits and isolation at the time of early range expansions into the region. The second is analogous to the genetic model of isolation by distance, and it predicts that genetic and linguistic correspondences formed through continuing genetic and linguistic exchange between neighboring populations. We tested the predictions of the two models by comparing observed and simulated patterns of genetic variation, genetic and linguistic trees, and matrices of genetic, linguistic, and geographic distances. The data consist of 751 autosomal microsatellites and 108 structural linguistic features collected from 33 Northern Island Melanesian populations. The results of the tests indicate that linguistic and genetic exchange have erased any evidence of a splitting and isolation process that might have occurred early in the settlement history of the region. The correlation patterns are also inconsistent with the predictions of the isolation by distance coevolutionary process in the larger Northern Island Melanesian region, but there is strong evidence for the process in the rugged interior of the largest island in the region (New Britain). There we found some of the strongest recorded correlations between genetic, linguistic, and geographic distances. We also found that, throughout the region, linguistic features have generally been less likely to diffuse across population boundaries than genes. The results from our study, based on exceptionally fine-grained data, show that local genetic and linguistic exchange are likely to obscure evidence of the early history of a region, and that language barriers do not particularly hinder genetic exchange. In contrast, global patterns may emphasize more ancient demographic events, including population splits associated with the early colonization of major world regions.     

Predictions of native American population structure using linguistic covariates in a hidden regression framework

Background

The mainland of the Americas is home to a remarkable diversity of languages, and the relationships between genes and languages have attracted considerable attention in the past. Here we investigate to which extent geography and languages can predict the genetic structure of Native American populations.

Methodology/Principal Findings

Our approach is based on a Bayesian latent cluster regression model in which cluster membership is explained by geographic and linguistic covariates. After correcting for geographic effects, we find that the inclusion of linguistic information improves the prediction of individual membership to genetic clusters. We further compare the predictive power of Greenberg''s and The Ethnologue classifications of Amerindian languages. We report that The Ethnologue classification provides a better genetic proxy than Greenberg''s classification at the stock and at the group levels. Although high predictive values can be achieved from The Ethnologue classification, we nevertheless emphasize that Choco, Chibchan and Tupi linguistic families do not exhibit a univocal correspondence with genetic clusters.

Conclusions/Significance

The Bayesian latent class regression model described here is efficient at predicting population genetic structure using geographic and linguistic information in Native American populations.     

Spatial congruence in language and species richness but not threat in the world's top linguistic hotspot

Languages share key evolutionary properties with biological species, and global-level spatial congruence in richness and threat is documented between languages and several taxonomic groups. However, there is little understanding of the functional connection between diversification or extinction in languages and species, or the relationship between linguistic and species richness across different spatial scales. New Guinea is the world''s most linguistically rich region and contains extremely high biological diversity. We demonstrate significant positive relationships between language and mammal richness in New Guinea across multiple spatial scales, revealing a likely functional relationship over scales at which infra-island diversification may occur. However, correlations are driven by spatial congruence between low levels of language and species richness. Regional biocultural richness may have showed closer congruence before New Guinea''s linguistic landscape was altered by Holocene demographic events. In contrast to global studies, we demonstrate a significant negative correlation across New Guinea between areas with high levels of threatened languages and threatened mammals, indicating that landscape-scale threats differ between these groups. Spatial resource prioritization to conserve biodiversity may not benefit threatened languages, and conservation policy must adopt a multi-faceted approach to protect biocultural diversity as a whole.     

Environmental paleomicrobiology: using DNA preserved in aquatic sediments to its full potential

In-depth knowledge about spatial and temporal variation in microbial diversity and function is needed for a better understanding of ecological and evolutionary responses to global change. In particular, the study of microbial ancient DNA preserved in sediment archives from lakes and oceans can help us to evaluate the responses of aquatic microbes in the past and make predictions about future biodiversity change in those ecosystems. Recent advances in molecular genetic methods applied to the analysis of historically deposited DNA in sediments have not only allowed the taxonomic identification of past aquatic microbial communities but also enabled tracing their evolution and adaptation to episodic disturbances and gradual environmental change. Nevertheless, some challenges remain for scientists to take full advantage of the rapidly developing field of paleo-genetics, including the limited ability to detect rare taxa and reconstruct complete genomes for evolutionary studies. Here, we provide a brief review of some of the recent advances in the field of environmental paleomicrobiology and discuss remaining challenges related to the application of molecular genetic methods to study microbial diversity, ecology, and evolution in sediment archives. We anticipate that, in the near future, environmental paleomicrobiology will shed new light on the processes of microbial genome evolution and microbial ecosystem responses to quaternary environmental changes at an unprecedented level of detail. This information can, for example, aid geological reconstructions of biogeochemical cycles and predict ecosystem responses to environmental perturbations, including in the context of human-induced global changes.     

The Proximate Phonological Unit of Chinese-English Bilinguals: Proficiency Matters

An essential step to create phonology according to the language production model by Levelt, Roelofs and Meyer is to assemble phonemes into a metrical frame. However, recently, it has been proposed that different languages may rely on different grain sizes of phonological units to construct phonology. For instance, it has been proposed that, instead of phonemes, Mandarin Chinese uses syllables and Japanese uses moras to fill the metrical frame. In this study, we used a masked priming-naming task to investigate how bilinguals assemble their phonology for each language when the two languages differ in grain size. Highly proficient Mandarin Chinese-English bilinguals showed a significant masked onset priming effect in English (L2), and a significant masked syllabic priming effect in Mandarin Chinese (L1). These results suggest that their proximate unit is phonemic in L2 (English), and that bilinguals may use different phonological units depending on the language that is being processed. Additionally, under some conditions, a significant sub-syllabic priming effect was observed even in Mandarin Chinese, which indicates that L2 phonology exerts influences on L1 target processing as a consequence of having a good command of English.     

From the origin of sex-determining factors to the evolution of sex-determining systems

Sex determination is typically classified as either genotypic or environmental. However, this dichotomy obscures the developmental origin and evolutionary modification of determinants of sex, and therefore hinders an understanding of the processes that generates diversity in sex-determining systems. Recent research on reptiles and fish emphasizes that sex determination is a multifactorial regulatory process that is best understood as a threshold dichotomy rather than as the result of genetically inherited triggers of development. Here we critically assess the relationship between the developmental origin of sex-determining factors and evolutionary transitions in sex-determining systems. Our perspective emphasizes the importance of both genetic and nongenetic causes in evolution of sex determination and may help to generate predictions with respect to the evolutionary patterns of sex-determining systems and the underlying diversity of developmental and genetic regulatory networks.     

Multilocus patterns of polymorphism and selection across the X chromosome of Caenorhabditis remanei

Natural selection and neutral processes such as demography, mutation, and gene conversion all contribute to patterns of polymorphism within genomes. Identifying the relative importance of these varied components in evolution provides the principal challenge for population genetics. To address this issue in the nematode Caenorhabditis remanei, I sampled nucleotide polymorphism at 40 loci across the X chromosome. The site-frequency spectrum for these loci provides no evidence for population size change, and one locus presents a candidate for linkage to a target of balancing selection. Selection for codon usage bias leads to the non-neutrality of synonymous sites, and despite its weak magnitude of effect (N(e)s approximately 0.1), is responsible for profound patterns of diversity and divergence in the C. remanei genome. Although gene conversion is evident for many loci, biased gene conversion is not identified as a significant evolutionary process in this sample. No consistent association is observed between synonymous-site diversity and linkage-disequilibrium-based estimators of the population recombination parameter, despite theoretical predictions about background selection or widespread genetic hitchhiking, but genetic map-based estimates of recombination are needed to rigorously test for a diversity-recombination relationship. Coalescent simulations also illustrate how a spurious correlation between diversity and linkage-disequilibrium-based estimators of recombination can occur, due in part to the presence of unbiased gene conversion. These results illustrate the influence that subtle natural selection can exert on polymorphism and divergence, in the form of codon usage bias, and demonstrate the potential of C. remanei for detecting natural selection from genomic scans of polymorphism.     

The impact of founder events on chromosomal variability in multiply mating species

In species with heterogametic males, the relative levels of X chromosome versus autosome diversity hold key information about the evolutionary forces at work in a population. It has been shown that population size changes alter the ratio of X linked to autosomal (X/A) variation, with population size reductions and recent bottlenecks leading to decreased X/A diversity ratios. Here we use theory and simulation to investigate a separate demographic effect-that of founder events involving multiply mated females-and find that it leads to much stronger reductions in X/A diversity ratios than are produced by simple population size changes. Investigating the potential of this process to account for sharply reduced X-linked diversity in European Drosophila melanogaster, we find that this model yields predictions that are compatible with the empirical data.     

Strategy Choice Mediates the Link between Auditory Processing and Spelling

Relations among linguistic auditory processing, nonlinguistic auditory processing, spelling ability, and spelling strategy choice were examined. Sixty-three undergraduate students completed measures of auditory processing (one involving distinguishing similar tones, one involving distinguishing similar phonemes, and one involving selecting appropriate spellings for individual phonemes). Participants also completed a modified version of a standardized spelling test, and a secondary spelling test with retrospective strategy reports. Once testing was completed, participants were divided into phonological versus nonphonological spellers on the basis of the number of words they spelled using phonological strategies only. Results indicated a) moderate to strong positive correlations among the different auditory processing tasks in terms of reaction time, but not accuracy levels, and b) weak to moderate positive correlations between measures of linguistic auditory processing (phoneme distinction and phoneme spelling choice in the presence of foils) and spelling ability for phonological spellers, but not for nonphonological spellers. These results suggest a possible explanation for past contradictory research on auditory processing and spelling, which has been divided in terms of whether or not disabled spellers seemed to have poorer auditory processing than did typically developing spellers, and suggest implications for teaching spelling to children with good versus poor auditory processing abilities.     

Germs, genomes and genealogies

Genetic diversity in pathogen species contains information about evolutionary and epidemiological processes, including the origins and history of disease, the nature of the selective forces acting on pathogen genes and the role of recombination in generating genetic novelty. Here, we review recent developments in these fields and compare the use of population genetic, or population-model based, approaches to phylogenetic, or population-model free, methodologies. We show how simple epidemiological models can be related to the ancestral, or coalescent, process underlying samples from pathogen species, enabling detailed inference about pathogen biology from patterns of molecular variation.     

TransNewGuinea.org: An Online Database of New Guinea Languages

The island of New Guinea has the world’s highest linguistic diversity, with more than 900 languages divided into at least 23 distinct language families. This diversity includes the world’s third largest language family: Trans-New Guinea. However, the region is one of the world’s least well studied, and primary data is scattered across a wide range of publications and more often then not hidden in unpublished “gray” literature. The lack of primary research data on the New Guinea languages has been a major impediment to our understanding of these languages, and the history of the peoples in New Guinea. TransNewGuinea.org aims to collect data about these languages and place them online in a consistent format. This database will enable future research into the New Guinea languages with both traditional comparative linguistic methods and novel cutting-edge computational techniques. The long-term aim is to shed light into the prehistory of the peoples of New Guinea, and to understand why there is such major diversity in their languages.