A global analysis of matches and mismatches between human genetic and linguistic histories.

Human history is written in both our genes and our languages. The extent to which our biological and linguistic histories are congruent has been the subject of considerable debate, with clear examples of both matches and mismatches. To disentangle the patterns of demographic and cultural transmission, we need a global systematic assessment of matches and mismatches. Here, we assemble a genomic database (GeLaTo, or Genes and Languages Together) specifically curated to investigate genetic and linguistic diversity worldwide. We find that most populations in GeLaTo that speak languages of the same language family (i.e., that descend from the same ancestor language) are also genetically highly similar. However, we also identify nearly 20% mismatches in populations genetically close to linguistically unrelated groups. These mismatches, which occur within the time depth of known linguistic relatedness up to about 10,000 y, are scattered around the world, suggesting that they are a regular outcome in human history. Most mismatches result from populations shifting to the language of a neighboring population that is genetically different because of independent demographic histories. In line with the regularity of such shifts, we find that only half of the language families in GeLaTo are genetically more cohesive than expected under spatial autocorrelations. Moreover, the genetic and linguistic divergence times of population pairs match only rarely, with Indo-European standing out as the family with most matches in our sample. Together, our database and findings pave the way for systematically disentangling demographic and cultural history and for quantifying processes of shifts in language and social identities on a global scale.

Neural signatures of syntactic variation in speech planning.

Planning to speak is a challenge for the brain, and the challenge varies between and within languages. Yet, little is known about how neural processes react to these variable challenges beyond the planning of individual words. Here, we examine how fundamental differences in syntax shape the time course of sentence planning. Most languages treat alike (i.e., align with each other) the 2 uses of a word like "gardener" in "the gardener crouched" and in "the gardener planted trees." A minority keeps these formally distinct by adding special marking in 1 case, and some languages display both aligned and nonaligned expressions. Exploiting such a contrast in Hindi, we used electroencephalography (EEG) and eye tracking to suggest that this difference is associated with distinct patterns of neural processing and gaze behavior during early planning stages, preceding phonological word form preparation. Planning sentences with aligned expressions induces larger synchronization in the theta frequency band, suggesting higher working memory engagement, and more visual attention to agents than planning nonaligned sentences, suggesting delayed commitment to the relational details of the event. Furthermore, plain, unmarked expressions are associated with larger desynchronization in the alpha band than expressions with special markers, suggesting more engagement in information processing to keep overlapping structures distinct during planning. Our findings contrast with the observation that the form of aligned expressions is simpler, and they suggest that the global preference for alignment is driven not by its neurophysiological effect on sentence planning but by other sources, possibly by aspects of production flexibility and fluency or by sentence comprehension. This challenges current theories on how production and comprehension may affect the evolution and distribution of syntactic variants in the world's languages.

Multi-variate coding for possession: methodology and preliminary results.

In this work we are presenting a database structure to encode the phenomenon of differential possession across languages, considering noun possession classes and possessive constructions as independent but linked. We show how this structure can be used to study different dimensions of possession: semantics, noun valence, and possessive constructions. We present preliminary survey results from a global sample of 120 languages and show that there is a universal semantic core in both inalienable and non-possessible noun classes. Inalienables are centered on body parts and kinship. Non-possessibles are centered on animals, humans, and natural elements.

Compositionality in animals and humans.

A key step in understanding the evolution of human language involves unravelling the origins of language's syntactic structure. One approach seeks to reduce the core of syntax in humans to a single principle of recursive combination, merge, for which there is no evidence in other species. We argue for an alternative approach. We review evidence that beneath the staggering complexity of human syntax, there is an extensive layer of nonproductive, nonhierarchical syntax that can be fruitfully compared to animal call combinations. This is the essential groundwork that must be explored and integrated before we can elucidate, with sufficient precision, what exactly made it possible for human language to explode its syntactic capacity, transitioning from simple nonproductive combinations to the unrivalled complexity that we now have.

Nouns slow down speech across structurally and culturally diverse languages.

By force of nature, every bit of spoken language is produced at a particular speed. However, this speed is not constant-speakers regularly speed up and slow down. Variation in speech rate is influenced by a complex combination of factors, including the frequency and predictability of words, their information status, and their position within an utterance. Here, we use speech rate as an index of word-planning effort and focus on the time window during which speakers prepare the production of words from the two major lexical classes, nouns and verbs. We show that, when naturalistic speech is sampled from languages all over the world, there is a robust cross-linguistic tendency for slower speech before nouns compared with verbs, both in terms of slower articulation and more pauses. We attribute this slowdown effect to the increased amount of planning that nouns require compared with verbs. Unlike verbs, nouns can typically only be used when they represent new or unexpected information; otherwise, they have to be replaced by pronouns or be omitted. These conditions on noun use appear to outweigh potential advantages stemming from differences in internal complexity between nouns and verbs. Our findings suggest that, beneath the staggering diversity of grammatical structures and cultural settings, there are robust universals of language processing that are intimately tied to how speakers manage referential information when they communicate with one another.

Dwarf mongoose alarm calls: investigating a complex non-human animal call.

Communication plays a vital role in the social lives of many species and varies greatly in complexity. One possible way to increase communicative complexity is by combining signals into longer sequences, which has been proposed as a mechanism allowing species with a limited repertoire to increase their communicative output. In mammals, most studies on combinatoriality have focused on vocal communication in non-human primates. Here, we investigated a potential combination of alarm calls in the dwarf mongoose (Helogale parvula), a non-primate mammal. Acoustic analyses and playback experiments with a wild population suggest: (i) that dwarf mongooses produce a complex call type (T3) which, at least at the surface level, seems to comprise units that are not functionally different to two meaningful alarm calls (aerial and terrestrial); and (ii) that this T3 call functions as a general alarm, produced in response to a wide range of threats. Using a novel approach, we further explored multiple interpretations of the T3 call based on the information content of the apparent comprising calls and how they are combined. We also considered an alternative, non-combinatorial interpretation that frames T3 as the origin, rather than the product, of the individual alarm calls. This study complements previous knowledge of vocal combinatoriality in non-primate mammals and introduces an approach that could facilitate comparisons between different animal and human communication systems.

Environmental factors drive language density more in food-producing than in hunter-gatherer populations.

Linguistic diversity is a key aspect of human population diversity and shapes much of our social and cognitive lives. To a considerable extent, the distribution of this diversity is driven by environmental factors such as climate or coast access. An unresolved question is whether the relevant factors have remained constant over time. Here, we address this question at a global scale. We approximate the difference between pre- versus post-Neolithic populations by the difference between modern hunter-gatherer versus food-producing populations. Using a novel geostatistical approach of estimating language and language family densities, we show that environmental-chiefly climate factors-have driven the language density of food-producing populations considerably more strongly than the language density of hunter-gatherer populations. Current evidence suggests that the population dynamics of modern hunter-gatherers is very similar to that of what can be reconstructed from the Palaeolithic record. Based on this, we cautiously infer that the impact of environmental factors on language densities underwent a substantial change with the transition to agriculture. After this transition, the environmental impact on language diversity in food-producing populations has remained relatively stable since it can also be detected-albeit in slightly weaker form-in models that capture the reduced linguistic diversity during large-scale language spreads in the Mid-Holocene.

Language evolution: syntax before phonology?

Phonology and syntax represent two layers of sound combination central to language's expressive power. Comparative animal studies represent one approach to understand the origins of these combinatorial layers. Traditionally, phonology, where meaningless sounds form words, has been considered a simpler combination than syntax, and thus should be more common in animals. A linguistically informed review of animal call sequences demonstrates that phonology in animal vocal systems is rare, whereas syntax is more widespread. In the light of this and the absence of phonology in some languages, we hypothesize that syntax, present in all languages, evolved before phonology.

Language follows a distinct mode of extra-genomic evolution.

As one of the most specific, yet most diverse of human behaviors, language is shaped by both genomic and extra-genomic evolution. Sharing methods and models between these modes of evolution has significantly advanced our understanding of language and inspired generalized theories of its evolution. Progress is hampered, however, by the fact that the extra-genomic evolution of languages, i.e. linguistic evolution, maps only partially to other forms of evolution. Contrasting it with the biological evolution of eukaryotes and the cultural evolution of technology as the best understood models, we show that linguistic evolution is special by yielding a stationary dynamic rather than stable solutions, and that this dynamic allows the use of language change for social differentiation while maintaining its global adaptiveness. Linguistic evolution furthermore differs from technological evolution by requiring vertical transmission, allowing the reconstruction of phylogenies; and it differs from eukaryotic biological evolution by foregoing a genotype vs phenotype distinction, allowing deliberate and biased change. Recognising these differences will improve our empirical tools and open new avenues for analyzing how linguistic, cultural, and biological evolution interacted with each other when language emerged in the hominin lineage. Importantly, our framework will help to cope with unprecedented scientific and ethical challenges that presently arise from how rapid cultural evolution impacts language, most urgently from interventional clinical tools for language disorders, potential epigenetic effects of technology on language, artificial intelligence and linguistic communicators, and global losses of linguistic diversity and identity. Beyond language, the distinctions made here allow identifying variation in other forms of biological and cultural evolution, developing new perspectives for empirical research.

A universal preference for animate agents in hominids.

When conversing, humans instantaneously predict meaning from fragmentary and ambiguous mspeech, long before utterance completion. They do this by integrating priors (initial assumptions about the world) with contextual evidence to rapidly decide on the most likely meaning. One powerful prior is attentional preference for agents, which biases sentence processing but universally so only if agents are animate. Here, we investigate the evolutionary origins of this preference, by allowing chimpanzees, gorillas, orangutans, human children, and adults to freely choose between agents and patients in still images, following video clips depicting their dyadic interaction. All participants preferred animate (and occasionally inanimate) agents, although the effect was attenuated if patients were also animate. The findings suggest that a preference for animate agents evolved before language and is not reducible to simple perceptual biases. To conclude, both humans and great apes prefer animate agents in decision tasks, echoing a universal prior in human language processing.

Surprisal From Language Models Can Predict ERPs in Processing Predicate-Argument Structures Only if Enriched by an Agent Preference Principle.

Language models based on artificial neural networks increasingly capture key aspects of how humans process sentences. Most notably, model-based surprisals predict event-related potentials such as N400 amplitudes during parsing. Assuming that these models represent realistic estimates of human linguistic experience, their success in modeling language processing raises the possibility that the human processing system relies on no other principles than the general architecture of language models and on sufficient linguistic input. Here, we test this hypothesis on N400 effects observed during the processing of verb-final sentences in German, Basque, and Hindi. By stacking Bayesian generalised additive models, we show that, in each language, N400 amplitudes and topographies in the region of the verb are best predicted when model-based surprisals are complemented by an Agent Preference principle that transiently interprets initial role-ambiguous noun phrases as agents, leading to reanalysis when this interpretation fails. Our findings demonstrate the need for this principle independently of usage frequencies and structural differences between languages. The principle has an unequal force, however. Compared to surprisal, its effect is weakest in German, stronger in Hindi, and still stronger in Basque. This gradient is correlated with the extent to which grammars allow unmarked NPs to be patients, a structural feature that boosts reanalysis effects. We conclude that language models gain more neurobiological plausibility by incorporating an Agent Preference. Conversely, theories of human processing profit from incorporating surprisal estimates in addition to principles like the Agent Preference, which arguably have distinct evolutionary roots.

Deep history of cultural and linguistic evolution among Central African hunter-gatherers.

Human evolutionary history in Central Africa reflects a deep history of population connectivity. However, Central African hunter-gatherers (CAHGs) currently speak languages acquired from their neighbouring farmers. Hence it remains unclear which aspects of CAHG cultural diversity results from long-term evolution preceding agriculture and which reflect borrowing from farmers. On the basis of musical instruments, foraging tools, specialized vocabulary and genome-wide data from ten CAHG populations, we reveal evidence of large-scale cultural interconnectivity among CAHGs before and after the Bantu expansion. We also show that the distribution of hunter-gatherer musical instruments correlates with the oldest genomic segments in our sample predating farming. Music-related words are widely shared between western and eastern groups and likely precede the borrowing of Bantu languages. In contrast, subsistence tools are less frequently exchanged and may result from adaptation to local ecologies. We conclude that CAHG material culture and specialized lexicon reflect a long evolutionary history in Central Africa.

An Agent-First Preference in a Patient-First Language During Sentence Comprehension.

The language comprehension system preferentially assumes that agents come first during incremental processing. While this might reflect a biologically fixed bias, shared with other domains and other species, the evidence is limited to languages that place agents first, and so the bias could also be learned from usage frequency. Here, we probe the bias with electroencephalography (EEG) in Äiwoo, a language that by default places patients first, but where sentence-initial nouns are still locally ambiguous between patient or agent roles. Comprehenders transiently interpreted nonhuman nouns as patients, eliciting a negativity when disambiguation was toward the less common agent-initial order. By contrast and against frequencies, human nouns were transiently interpreted as agents, eliciting an N400-like negativity when the disambiguation was toward patient-initial order. Consistent with the notion of a fixed property, the agent bias is robust against usage frequency for human referents. However, this bias can be reversed by frequency experience for nonhuman referents.

The Agent Preference in Visual Event Apprehension.

A central aspect of human experience and communication is understanding events in terms of agent ("doer") and patient ("undergoer" of action) roles. These event roles are rooted in general cognition and prominently encoded in language, with agents appearing as more salient and preferred over patients. An unresolved question is whether this preference for agents already operates during apprehension, that is, the earliest stage of event processing, and if so, whether the effect persists across different animacy configurations and task demands. Here we contrast event apprehension in two tasks and two languages that encode agents differently; Basque, a language that explicitly case-marks agents ('ergative'), and Spanish, which does not mark agents. In two brief exposure experiments, native Basque and Spanish speakers saw pictures for only 300 ms, and subsequently described them or answered probe questions about them. We compared eye fixations and behavioral correlates of event role extraction with Bayesian regression. Agents received more attention and were recognized better across languages and tasks. At the same time, language and task demands affected the attention to agents. Our findings show that a general preference for agents exists in event apprehension, but it can be modulated by task and language demands.

Nouns and verbs in Chintang: children's usage and surrounding adult speech.

Analyzing the development of the noun-to-verb ratio in a longitudinal corpus of four Chintang (Sino-Tibetan) children, we find that up to about age four, children have a significantly higher ratio than adults. Previous cross-linguistic research rules out an explanation of this in terms of a universal noun bias; instead, a likely cause is that Chintang verb morphology is polysynthetic and difficult to learn. This hypothesis is supported by the fact that the development of Chintang children's noun-to-verb ratio correlates significantly with the extent to which they show a similar flexibility with verbal morphology to that of the surrounding adults, as measured by morphological paradigm entropy. While this development levels off around age three, children continue to have a higher overall noun-to-verb ratio than adults. A likely explanation lies in the kinds of activities that children are engaged in and that are almost completely separate from adults' activities in this culture.

Transition to language: From agent perception to event representation.

Spoken language, as we have it, requires specific capacities-at its most basic advanced vocal control and complex social cognition. In humans, vocal control is the basis for speech, achieved through coordinated interactions of larynx activity and rapid changes in vocal tract configurations. Most likely, speech evolved in response to early humans perceiving reality in increasingly complex ways, to the effect that primate-like signaling became unsustainable as a sole communication device. However, in what ways did and do humans see the world in more complex ways compared to other species? Although animal signals can refer to external events, in contrast to humans, they usually refer to the agents only, sometimes in compositional ways, but never together with patients. It may be difficult for animals to comprehend events as part of larger social scripts, with antecedent causes and future consequences, which are more typically tie the patient into the event. Human brain enlargement over the last million years probably has provided the cognitive resources to represent social interactions as part of bigger social scripts, which enabled humans to go beyond an agent-focus to refer to agent-patient relations, the likely foundation for the evolution of grammar. This article is categorized under: Cognitive Biology > Evolutionary Roots of Cognition Linguistics > Evolution of Language Psychology > Comparative.

More or Less Unnatural: Semantic Similarity Shapes the Learnability and Cross-Linguistic Distribution of Unnatural Syncretism in Morphological Paradigms.

Morphological systems often reuse the same forms in different functions, creating what is known as syncretism. While syncretism varies greatly, certain cross-linguistic tendencies are apparent. Patterns where all syncretic forms share a morphological feature value (e.g., first person, or plural number) are most common cross-linguistically, and this preference is mirrored in results from learning experiments. While this suggests a general bias towards natural (featurally homogeneous) over unnatural (featurally heterogeneous) patterns, little is yet known about gradients in learnability and distributions of different kinds of unnatural patterns. In this paper we assess apparent cross-linguistic asymmetries between different types of unnatural patterns in person-number verbal agreement paradigms and test their learnability in an artificial language learning experiment. We find that the cross-linguistic recurrence of unnatural patterns of syncretism in person-number paradigms is proportional to the amount of shared feature values (i.e., semantic similarity) amongst the syncretic forms. Our experimental results further suggest that the learnability of syncretic patterns also mirrors the paradigm's degree of feature-value similarity. We propose that this gradient in learnability reflects a general bias towards similarity-based structure in morphological learning, which previous literature has shown to play a crucial role in word learning as well as in category and concept learning more generally. Rather than a dichotomous natural/unnatural distinction, our results thus support a more nuanced view of (un)naturalness in morphological paradigms and suggest that a preference for similarity-based structure during language learning might shape the worldwide transmission and typological distribution of patterns of syncretism.

The evolutionary origins of syntax: Event cognition in nonhuman primates.

Languages tend to encode events from the perspective of agents, placing them first and in simpler forms than patients. This agent bias is mirrored by cognition: Agents are more quickly recognized than patients and generally attract more attention. This leads to the hypothesis that key aspects of language structure are fundamentally rooted in a cognition that decomposes events into agents, actions, and patients, privileging agents. Although this type of event representation is almost certainly universal across languages, it remains unclear whether the underlying cognition is uniquely human or more widespread in animals. Here, we review a range of evidence from primates and other animals, which suggests that agent-based event decomposition is phylogenetically older than humans. We propose a research program to test this hypothesis in great apes and human infants, with the goal to resolve one of the major questions in the evolution of language, the origins of syntax.

From collocations to call-ocations: using linguistic methods to quantify animal call combinations.

Abstract: Emerging data in a range of non-human animal species have highlighted a latent ability to combine certain pre-existing calls together into larger structures. Currently, however, the quantification of context-specific call combinations has received less attention. This is problematic because animal calls can co-occur with one another simply through chance alone. One common approach applied in language sciences to identify recurrent word combinations is collocation analysis. Through comparing the co-occurrence of two words with how each word combines with other words within a corpus, collocation analysis can highlight above chance, two-word combinations. Here, we demonstrate how this approach can also be applied to non-human animal signal sequences by implementing it on artificially generated data sets of call combinations. We argue collocation analysis represents a promising tool for identifying non-random, communicatively relevant call combinations and, more generally, signal sequences, in animals. Significance statement: Assessing the propensity for animals to combine calls provides important comparative insights into the complexity of animal vocal systems and the selective pressures such systems have been exposed to. Currently, however, the objective quantification of context-specific call combinations has received less attention. Here we introduce an approach commonly applied in corpus linguistics, namely collocation analysis, and show how this method can be put to use for identifying call combinations more systematically. Through implementing the same objective method, so-called call-ocations, we hope researchers will be able to make more meaningful comparisons regarding animal signal sequencing abilities both within and across systems. Supplementary Information: The online version contains supplementary material available at 10.1007/s00265-022-03224-3.

Cross-linguistic differences in case marking shape neural power dynamics and gaze behavior during sentence planning.

Languages differ in how they mark the dependencies between verbs and arguments, e.g., by case. An eye tracking and EEG picture description study examined the influence of case marking on the time course of sentence planning in Basque and Swiss German. While German assigns an unmarked (nominative) case to subjects, Basque specifically marks agent arguments through ergative case. Fixations to agents and event-related synchronization (ERS) in the theta and alpha frequency bands, as well as desynchronization (ERD) in the alpha and beta bands revealed multiple effects of case marking on the time course of early sentence planning. Speakers decided on case marking under planning early when preparing sentences with ergative-marked agents in Basque, whereas sentences with unmarked agents allowed delaying structural commitment across languages. These findings support hierarchically incremental accounts of sentence planning and highlight how cross-linguistic differences shape the neural dynamics underpinning language use.