Publications

Abstract

Computer models of cultural evolution have shown language properties emerging on interacting agents with a brain that lacks dedicated, nativist language modules. Notably, models using Bayesian agents provide a precise specification of (extra-)liguististic factors (e.g., genetic) that shape language through iterated learning (biases on language), and demonstrate that weak biases get expressed more strongly over time (bias amplification). Other models attempt to lessen assumption on agents’ innate predispositions even more, and emphasize self-organization within agents, highlighting glossogenesis (the development of language from a nonlinguistic state). Ultimately however, one also has to recognize that biology and culture are strongly interacting, forming a coevolving system. As such, computer models show that agents might (biologically) evolve to a state predisposed to language adaptability, where (culturally) stable language features might get assimilated into the genome via Baldwinian niche construction. In summary, while many questions about language evolution remain unanswered, it is clear that it is not to be completely understood from a purely biological, cognitivist perspective. Language should be regarded as (partially) emerging on the social interactions between large populations of speakers. In this context, agent models provide a sound approach to investigate the complex dynamics of genetic biasing on language and speech

Abstract

People vary at most levels, from the molecular to the cognitive, and the shape of the hard palate (the bony roof of the mouth) is no exception. The patterns of variation in the hard palate are important for the forensic sciences and (palaeo)anthropology, and might also play a role in speech production, both in pathological cases and normal variation. Here we describe a method based on Bézier curves, whose main aim is to generate possible shapes of the hard palate in humans for use in computer simulations of speech production and language evolution. Moreover, our method can also capture existing patterns of variation using few and easy-to-interpret parameters, and fits actual data obtained from MRI traces very well with as little as two or three free parameters. When compared to the widely-used Principal Component Analysis (PCA), our method fits actual data slightly worse for the same number of degrees of freedom. However, it is much better at generating new shapes without requiring a calibration sample, its parameters have clearer interpretations, and their ranges are grounded in geometrical considerations. © 2018 Janssen et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Abstract

Children's abilities to process the phonological structure of words are important predictors of their literacy development. In the current study, we examined the interrelatedness between implicit (i.e., speech decoding) and explicit (i.e., phonological awareness) phonological abilities, and especially the role therein of lexical specificity (i.e., the ability to learn to recognize spoken words based on only minimal acoustic-phonetic differences). We tested 75 Dutch monolingual and 64 Turkish–Dutch bilingual kindergartners. SEM analyses showed that speech decoding predicted lexical specificity, which in turn predicted rhyme awareness in the first language learners but phoneme awareness in the second language learners. Moreover, in the latter group there was an impact of the second language: Dutch speech decoding and lexical specificity predicted Turkish phonological awareness, which in turn predicted Dutch phonological awareness. We conclude that language-specific phonological characteristics underlie different patterns of transfer from implicit to explicit phonological abilities in first and second language learners.

Abstract

Talkers are recognized more accurately if they are speaking the listeners’ native language rather than an unfamiliar language. This “language familiarity effect” has been shown not to depend upon comprehension and must instead involve language sound patterns. We further examine the level of sound-pattern processing involved, by comparing talker recognition in foreign languages versus two varieties of English, by (a) English speakers of one variety, (b) English speakers of the other variety, and (c) non-native listeners (more familiar with one of the varieties). All listener groups performed better with native than foreign speech, but no effect of language variety appeared: Native listeners discriminated talkers equally well in each, with the native variety never outdoing the other variety, and non-native listeners discriminated talkers equally poorly in each, irrespective of the variety's familiarity. The results suggest that this talker recognition effect rests not on simple familiarity, but on an abstract level of phonological processing

Abstract

Children who hear large amounts of diverse speech learn language more quickly than children who do not. However, high correlations between the amount and the diversity of the input in speech samples makes it difficult to isolate the influence of each. We overcame this problem by controlling the input to a computational model so that amount of exposure to linguistic input (quantity) and the quality of that input (lexical diversity) were independently manipulated. Sublexical, lexical, and multi-word knowledge were charted across development (Study 1), showing that while input quantity may be important early in learning, lexical diversity is ultimately more crucial, a prediction confirmed against children’s data (Study 2). The model trained on a lexically diverse input also performed better on nonword repetition and sentence recall tests (Study 3) and was quicker to learn new words over time (Study 4). A language input that is rich in lexical diversity outperforms equivalent richness in quantity for learned sublexical and lexical knowledge, for well-established language tests, and for acquiring words that have never been encountered before.

Abstract

Sustained attention has previously been shown as a requirement for language production. However, this is mostly evident for difficult conditions, such as a dual-task situation. The current study provides corroborating evidence that this relationship holds even for simple picture naming. Sustained attention ability, indexed both by participants’ reaction times and individuals’ hit rate (the proportion of correctly detected targets) on a digit discrimination task, correlated with picture naming latencies. Individuals with poor sustained attention were consistently slower and their RT distributions were more positively skewed when naming pictures compared to individuals with better sustained attention. Additionally, the need to sustain attention was manipulated by changing the speed of stimulus presentation. Research has suggested that fast event rates tax sustained attention resources to a larger degree than slow event rates. However, in this study the fast event rate did not result in increased difficulty, neither for the picture naming task nor for the sustained attention task. Instead, the results point to a speed-accuracy trade-off in the sustained attention task (lower accuracy but faster responses in the fast than in the slow event rate), and to a benefit for faster rates in the picture naming task (shorter naming latencies with no difference in accuracy). Performance on both tasks was largely comparable, supporting previous findings that sustained attention is called upon during language production

Abstract

Children with specific language impairment (SLI) have problems not only with language performance but also with sustained attention, which is the ability to maintain alertness over an extended period of time. Although there is consensus that this ability is impaired with respect to processing stimuli in the auditory perceptual modality, conflicting evidence exists concerning the visual modality.
Aims

To address the outstanding issue whether the impairment in sustained attention is limited to the auditory domain, or if it is domain-general. Furthermore, to test whether children's sustained attention ability relates to their word-production skills.
Methods & Procedures

Groups of 7–9 year olds with SLI (N = 28) and typically developing (TD) children (N = 22) performed a picture-naming task and two sustained attention tasks, namely auditory and visual continuous performance tasks (CPTs).
Outcomes & Results

Children with SLI performed worse than TD children on picture naming and on both the auditory and visual CPTs. Moreover, performance on both the CPTs correlated with picture-naming latencies across developmental groups.
Conclusions & Implications

These results provide evidence for a deficit in both auditory and visual sustained attention in children with SLI. Moreover, the study indicates there is a relationship between domain-general sustained attention and picture-naming performance in both TD and language-impaired children. Future studies should establish whether this relationship is causal. If attention influences language, training of sustained attention may improve language production in children from both developmental groups.

Abstract

Theories of conversation propose that in order to have smooth transitions from one turn to the next, speakers already plan their response while listening to their interlocutor. Moreover, it has been argued that speakers align their linguistic representations (i.e. prime each other), thereby reducing the processing costs associated with concurrent listening and speaking. In two experiments, we assessed how identity and associative priming from spoken words onto picture naming were affected by a concurrent speech planning task. In a baseline (no name) condition, participants heard prime words that were identical, associatively related, or unrelated to target pictures presented two seconds after prime onset. Each prime was accompanied by a non-target picture and followed by its recorded name. The participant did not name the non-target picture. In the plan condition, the participants first named the non-target picture, instead of listening to the recording, and then the target. In Experiment 1, where the plan- and no-plan conditions were tested between participants, priming effects of equal strength were found in the plan and no-plan condition. In Experiment 2, where the two conditions were tested within participants, the identity priming effect was maintained, but the associative priming effect was only seen in the no-plan but not in the plan condition. In this experiment, participant had to decide at the onset of each trial whether or not to name the non-target picture, rendering the task more complex than in Experiment 1. These decision processes may have interfered with the processing of the primes. Thus, associative priming can take place during speech planning, but only if the cognitive load is not too high.

Abstract

Spontaneous language learning both in children learning their mother tongue and in adults learning a second language shows that language development proceeds in a stage-wise manner. Given that a developmental stage is defined as a coherent linguistic system, utterances of language learners can be accounted for in terms of what (Selinker, Larry. 1972. Interlanguage. International Review of Applied Linguistics 10. 209-231) referred to with the term Interlanguage. This paper is a study on the early interlanguage systems of children learning Dutch and German as their mother tongue. The present child learner systems, so it is claimed, are coherent lexical systems based on types of verb-argument structure that are either agentive (as in Dutch: kannie bal pakke 'cannot ball get', or German: mag nich nase putzen 'like not nose clean') or non-agentive (as in Dutch: popje valt bijna 'doll falls nearly', or in German: ente fällt 'duck falls'). At this lexical stage, functional morphology (e. g. morphological finiteness, tense), function words (e. g. auxiliary verbs, determiners) and word order variation are absent. For these typically developing children, both in Dutch and in German, it is claimed that developmental progress is driven by the acquisition of the formal properties of topicalization. It is, furthermore, argued that this feature seems to serve as the driving force in the instantiation of the functional, i. e. informational linguistic properties of the target-language system

Abstract

The mutual exclusivity (ME) assumption is proposed to facilitate early word learning by guiding infants to map novel words to novel referents. This study assessed the emergence and use of ME to both disambiguate and retain the meanings of novel words across development in 18‐month‐old monolingual and bilingual children (Experiment 1; N = 58), and in a sub‐group of these children again at 24 months of age (Experiment 2: N = 32). Both monolinguals and bilinguals employed ME to select the referent of a novel label to a similar extent at 18 and 24 months. At 18 months, there were also no differences in novel word retention between the two language‐background groups. However, at 24 months, only monolinguals showed the ability to retain these label–object mappings. These findings indicate that the development of the ME assumption as a reliable word‐learning strategy is shaped by children's individual language exposure and experience with language use.

Abstract

In this article, we review research on child–robot interaction (CRI) to discuss how social robots can be used to scaffold language learning in young children. First we provide reasons why robots can be useful for teaching first and second languages to children. Then we review studies on CRI that used robots to help children learn vocabulary and produce language. The studies vary in first and second languages and demographics of the learners (typically developing children and children with hearing and communication impairments). We conclude that, although social robots are useful for teaching language to children, evidence suggests that robots are not as effective as human teachers. However, this conclusion is not definitive because robots that tutor students in language have not been evaluated rigorously and technology is advancing rapidly. We suggest that CRI offers an opportunity for research and list possible directions for that work.

Abstract

It has been suggested that some individuals may benefit more from social robots than do others. Using second
language (L2) as an example, the present study examined how individual differences in attitudes toward robots and personality
traits may be related to learning outcomes. Preliminary results with 24 Turkish-speaking adults suggest that negative attitudes
toward robots, more specifically thoughts and anxiety about the negative social impact that robots may have on the society,
predicted how well adults learned L2 words from a social robot. The possible implications of the findings as well as future directions are also discussed

Abstract

In production, frequently used words are preferentially extended to new, though related meanings. In comprehension, frequent exposure to a word instead makes the learner confident that all of the word’s legitimate uses have been experienced, resulting in an entrenched form-meaning mapping between the word and its experienced meaning(s). This results in a perception-production dissociation, where the forms speakers are most likely to map onto a novel meaning are precisely the forms that they believe can never be used that way. At first glance, this result challenges the idea of bidirectional form-meaning mappings, assumed by all current approaches to linguistic theory. In this paper, we show that bidirectional form-meaning mappings are not in fact challenged by this production-perception dissociation. We show that the production-perception dissociation is expected even if learners of the lexicon acquire simple symmetrical form-meaning associations through simple Hebbian learning.

Abstract

Deaf children born to hearing parents are exposed to language input quite late, which has long-lasting effects on language production. Previous studies with deaf individuals mostly focused on linguistic expressions of motion events, which have several event components. We do not know if similar effects emerge in simple events such as descriptions of spatial configurations of objects. Moreover, previous data mainly come from late adult signers. There is not much known about language development of late signing children soon after learning sign language. We compared simple event descriptions of late signers of Turkish Sign Language (adults, children) to age-matched native signers. Our results indicate that while late signers in both age groups are native-like in frequency of expressing a relational encoding, they lag behind native signers in using morphologically complex linguistic forms compared to other simple forms. Late signing children perform similar to adults and thus showed no development over time.

Abstract

There is evidence that the human cerebellum is involved not only in motor control but also in other cognitive functions. Several studies have shown that language-related activation is lateralized toward the right cerebellar hemisphere in most people, in accordance with leftward cerebral cortical lateralization for language and a general contralaterality of cerebral–cerebellar activations. In terms of behavior, hand use elicits asymmetrical activation in the cerebellum, while hand preference is weakly associated with language lateralization. However, it is not known how, or whether, these functional relations are reflected in anatomy. We investigated volumetric gray matter asymmetries of cerebellar lobules in an MRI data set comprising 2226 subjects. We tested these cerebellar asymmetries for associations with handedness, and for correlations with cerebral cortical anatomical asymmetries of regions important for language or hand motor control, as defined by two different automated image analysis methods and brain atlases, and supplemented with extensive visual quality control. No significant associations of cerebellar asymmetries to handedness were found. Some significant associations of cerebellar lobular asymmetries to cerebral cortical asymmetries were found, but none of these correlations were greater than 0.14, and they were mostly method-/atlas-dependent. On the basis of this large and highly powered study, we conclude that there is no overt structural manifestation of cerebellar functional lateralization and connectivity, in respect of hand motor control or language laterality

Abstract

English and German have similar prosody, but their speakers realize some pitch falls (not rises) in subtly different ways. We here test for asymmetry in perception. An ABX discrimination task requiring F0 slope or duration judgements on isolated vowels revealed no cross-language difference in duration or F0 fall discrimination, but discrimination of rises (realized similarly in each language) was less accurate for English than for German listeners. This unexpected finding may reflect greater sensitivity to rising patterns by German listeners, or reduced sensitivity by English listeners as a result of extensive exposure to phrase-final rises (“uptalk”) in their language

Abstract

In Dutch and German, the canonical order of subject, object(s) and finite verb is ‘verb-second’ (V2) in main but ‘verb-final’ (VF) in subordinate clauses. This occasionally leads to the production of noncanonical word orders. Familiar examples are causative and argumentative clauses introduced by a subordinating conjunction (Du. omdat, Ger. weil ‘because’): the omdat/weil-V2 phenomenon. Such clauses may also be introduced by coordinating conjunctions (Du. want, Ger. denn), which license V2 exclusively. However, want/denn-VF structures are unknown. We present the results of a corpus study on the incidence of omdat-V2 in spoken Dutch, and compare them to published data on weil-V2 in spoken German. Basic findings: omdat-V2 is much less frequent than weil-V2 (ratio almost 1:8); and the frequency relations between coordinating and subordinating conjunctions are opposite (want >> omdat; denn << weil). We propose that conjunction selection and V2/VF selection proceed partly independently, and sometimes miscommunicate—e.g. yielding omdat/weil paired with V2. Want/denn-VF pairs do not occur because want/denn clauses are planned as autonomous sentences, which take V2 by default. We sketch a simple feedforward neural network with two layers of nodes (representing conjunctions and word orders, respectively) that can simulate the observed data pattern through inhibition-based competition of the alternative choices within the node layers.

Abstract

In a paper entitled “Against markedness (and what to replace it with)”, Haspelmath argues “that the term ‘markedness’ is superfluous”, and that frequency asymmetries often explain structural (un)markedness asymmetries (Haspelmath 2006). We investigate whether this argument applies to Object and Verb orders in main (VO, marked) and subordinate (OV, unmarked) clauses of spoken and written German and Dutch, using English (without VO/OV alternation) as control. Frequency counts from six treebanks (three languages, two output modalities) do not support Haspelmath’s proposal. However, they reveal an unexpected phenomenon, most prominently in spoken Dutch and German: a small set of extremely high-frequent finite verbs with unspecific meanings populates main clauses much more densely than subordinate clauses. We suggest these verbs accelerate the start-up of grammatical encoding, thus facilitating sentence-initial output fluency

Abstract

Essays van psychologen en linguı̈sten over de relatie hersens-computers.

Abstract

A series of experiments is reported in which subjects describe simple visual scenes by means of both sentential and non-sentential responses. The data support the following statements about the lexicalization (word finding) process. (1) Words used by speakers in overt naming or sentence production responses are selected by a sequence of two lexical retrieval processes, the first yielding abstract pre-phonological items (Ll -items), the second one adding their phonological shapes (L2-items). (2) The selection of several Ll-items for a multi-word utterance can take place simultaneously. (3) A monitoring process is watching the output of Ll-lexicalization to check if it is in keeping with prevailing constraints upon utterance format. (4) Retrieval of the L2-item which corresponds with a given LI-item waits until the Ld-item has been checked by the monitor, and all other Ll-items needed for the utterance under construction have become available. A coherent picture of the lexicalization process begins to emerge when these characteristics are brought together with other empirical results in the area of naming and sentence production, e.g., picture naming reaction times (Seymour, 1979), speech errors (Garrett, 1980), and word order preferences (Bock, 1982).

Abstract

In this article, we examine gaze direction in responses to polar questions using both quantitative and conversation analytic (CA) methods. The data come from a novel corpus of conversations in which participants wore eye-tracking glasses to obtain direct measures of their eye movements. The results show that while most preferred responses are produced with gaze toward the questioner, most dispreferred responses are produced with gaze aversion. We further demonstrate that gaze aversion by respondents can occasion self-repair by questioners in the transition space between turns, indicating that the relationship between gaze direction and preference is more than a mere statistical association. We conclude that gaze direction in responses to polar questions functions as a signal of response preference. Data are in American, British, and Canadian English.

Abstract

Language development is driven by multiple factors involving both the individual child and the environments that surround the child. The chapters in this volume highlight several such factors as potential contributors to developmental change, including factors that examine the role of immediate social environment (i.e., parent SES, parent and sibling input, peer interaction) and factors that focus on the child’s own cognitive and social development, such as the acquisition of theory of mind, event knowledge, and memory. The discussion of the different factors is presented largely from a crosslinguistic framework, using a multimodal perspective (speech, gesture, sign). The book celebrates the scholarly contributions of Prof. Ayhan Aksu-Koç – a pioneer in the study of crosslinguistic variation in language acquisition, particularly in the domain of evidentiality and theory of mind. This book will serve as an important resource for researchers in the field of developmental psychology, cognitive science, and linguistics across the globe

Abstract

We report a large‐scale electrophysiological study of infant speech segmentation, in which over 100 English‐acquiring 9‐month‐olds were exposed to unfamiliar bisyllabic words embedded in sentences (e.g., He saw a wild eagle up there), after which their brain responses to either the just‐familiarized word (eagle) or a control word (coral) were recorded. When initial exposure occurs in continuous speech, as here, past studies have reported that even somewhat older infants do not reliably recognize target words, but that successful segmentation varies across children. Here, we both confirm and further uncover the nature of this variation. The segmentation response systematically varied across individuals and was related to their vocabulary development. About one‐third of the group showed a left‐frontally located relative negativity in response to familiar versus control targets, which has previously been described as a mature response. Another third showed a similarly located positive‐going reaction (a previously described immature response), and the remaining third formed an intermediate grouping that was primarily characterized by an initial response delay. A fine‐grained group‐level analysis suggested that a developmental shift to a lexical mode of processing occurs toward the end of the first year, with variation across individual infants in the exact timing of this shift.

Abstract

Humans differ in innumerable ways, with considerable variation observable at every level of description, from the molecular to the social. Traditionally, linguistic and psycholinguistic theory has downplayed the possibility of meaningful differences in language across individuals. However, it is becoming increasingly evident that there is
significant variation among speakers at any age as well as across the lifespan. In this paper, we review recent research in psycholinguistics, and argue that a focus on individual differences provides a crucial source of evidence that bears strongly upon core issues in theories of the acquisition and processing of language; specifically, the role of experience in language acquisition, processing, and attainment, and the architecture of the language faculty.

Abstract

We report three studies (one corpus, two experimental) that investigated the acquisition of relative clauses (RCs) in Finnish-speaking children. Study 1 found that Finnish children's naturalistic exposure to RCs predominantly consists of non-subject relatives (i.e. oblique, object) which typically have inanimate head nouns. Study 2 tested children's comprehension of subject, object, and two types of oblique relatives. No difference was found in the children's performance on different structures, including a lack of previously widely reported asymmetry between subject and object relatives. However, children's comprehension was modulated by animacy of the head referent. Study 3 tested children's production of the same RC structures using sentence repetition. Again we found no subject–object asymmetry. The pattern of results suggested that distributional frequency patterns and the relative complexity of the relativizer contribute to the difficulty associated with particular RC structures.

Abstract

Pretence and creativity are often regarded as ubiquitous characteristics of childhood, yet not all education systems value or promote these attributes to the same extent. Different pedagogies and practices are evident within the UK National Curriculum, Steiner and Montessori schools. In this study, 20 children participated from each of these school systems (N = 60, aged 6;10–8;11) completing the test of creative thinking—drawing production (TCT-DP; K. K. Urban, & H. G. Jellen, 1996) and a pretend actions task (W. F. Overton & J. P. Jackson, 1973). Overall, Steiner pupils performed significantly higher on the TCT-DP than both the Montessori and National Curriculum pupils who performed similarly. Steiner pupils also performed significantly better on the pretend actions task than the Montessori pupils, but no other significant differences were found. Overall, there was also a significant positive correlation between pretence and creativity in the current sample, supporting previous research suggesting that these skills are related (e.g., A.S. Kaugars & S. W. Russ, 2009; P. Y. Mullineaux & L. F. Dilalla, 2009).

Abstract

People spontaneously produce gestures during speaking and thinking. The authors focus here on gestures that depict or indicate information related to the contents of concurrent speech or thought (i.e., representational gestures). Previous research indicates that such gestures have not only communicative functions, but also self-oriented cognitive functions. In this article, the authors propose a new theoretical framework, the gesture-for-conceptualization hypothesis, which explains the self-oriented functions of representational gestures. According to this framework, representational gestures affect cognitive processes in 4 main ways: gestures activate, manipulate, package, and explore spatio-motoric information for speaking and thinking. These four functions are shaped by gesture's ability to schematize information, that is, to focus on a small subset of available information that is potentially relevant to the task at hand. The framework is based on the assumption that gestures are generated from the same system that generates practical actions, such as object manipulation; however, gestures are distinct from practical actions in that they represent information. The framework provides a novel, parsimonious, and comprehensive account of the self-oriented functions of gestures. The authors discuss how the framework accounts for gestures that depict abstract or metaphoric content, and they consider implications for the relations between self-oriented and communicative functions of gestures

Abstract

Sentence-final particles (SFPs) as bound morphemes in Japanese have no obvious effect on the truth conditions of a sentence. However, they encompass a diverse range of usages, from typical to atypical, according to the context and the interpersonal relationships in the specific situation. The most frequent particle,-ne, is typically used after addressee-oriented propositions for information sharing, while another frequent particle,-yo, is typically used after addresser-oriented propositions to elicit a sense of strength. This study sheds light on individual differences among native speakers in flexibly understanding such linguistic markers based on their mentalizing ability (i.e., the ability to infer the mental states of others). Two experiments employing electroencephalography (EEG) consistently showed enhanced early posterior negativities (EPN) for atypical SFP usage compared to typical usage, especially when understanding-ne compared to -yo, in both an SFP appropriateness judgment task and a content comprehension task. Importantly, the amplitude of the EPN for atypical usages of-ne was significantly higher in participants with lower mentalizing ability than in those with a higher mentalizing ability. This effect plausibly reflects low-ability mentalizers' stronger sense of strangeness toward atypical-ne usage. While high-ability mentalizers may aptly perceive others' attitudes via their various usages of-ne, low-ability mentalizers seem to adopt a more stereotypical understanding. These results attest to the greater degree of difficulty low-ability mentalizers have in establishing a smooth regulation of interpersonal distance during social encounters.

Abstract

The geographical region of Insular South East Asia and New Guinea is well-known as an
area of mega-biodiversity. Less well-known is the extreme linguistic diversity in this area:
over a quarter of the world’s 6,000 languages are spoken here. As small minority languages,
most of them will cease to be spoken in the coming few generations. The project described
here ensures the preservation of unique records of languages and the cultures encapsulated
by them in the region. The language resources were gathered by twenty linguists at,
or in collaboration with, Dutch universities over the last 40 years, and were compiled and
archived in collaboration with The Language Archive (TLA) at the Max Planck Institute in
Nijmegen. The resulting archive constitutes a collection ofmultimediamaterials and written
documents from 48 languages in Insular South East Asia and West New Guinea. At TLA,
the data was archived according to state-of-the-art standards (TLA holds the Data Seal of
Approval): the component metadata infrastructure CMDI was used; all metadata categories
as well as relevant units of annotation were linked to the ISO data category registry ISOcat.
This guaranteed proper integration of the language resources into the CLARIN framework.
Through the archive, future speaker communities and researchers will be able to extensively
search thematerials for answers to their own questions, even if they do not themselves know the language, and even if the language dies.

Abstract

Neurodevelopmental disorders are defined by highly heritable problems during development and brain growth. Attention-deficit/hyperactivity disorder (ADHD), autism spectrum disorders (ASDs), and intellectual disability (ID) are frequent neurodevelopmental disorders, with common comorbidity among them. Imaging genetics studies on the role of disease-linked genetic variants on brain structure and function have been performed to unravel the etiology of these disorders. Here, we reviewed imaging genetics literature on these disorders attempting to understand the mechanisms of individual disorders and their clinical overlap. For ADHD and ASD, we selected replicated candidate genes implicated through common genetic variants. For ID, which is mainly caused by rare variants, we included genes for relatively frequent forms of ID occurring comorbid with ADHD or ASD. We reviewed case-control studies and studies of risk variants in healthy individuals. Imaging genetics studies for ADHD were retrieved for SLC6A3/DAT1, DRD2, DRD4, NOS1, and SLC6A4/5HTT. For ASD, studies on CNTNAP2, MET, OXTR, and SLC6A4/5HTT were found. For ID, we reviewed the genes FMR1, TSC1 and TSC2, NF1, and MECP2. Alterations in brain volume, activity, and connectivity were observed. Several findings were consistent across studies, implicating, for example, SLC6A4/5HTT in brain activation and functional connectivity related to emotion regulation. However, many studies had small sample sizes, and hypothesis-based, brain region-specific studies were common. Results from available studies confirm that imaging genetics can provide insight into the link between genes, disease-related behavior, and the brain. However, the field is still in its early stages, and conclusions about shared mechanisms cannot yet be drawn.

Abstract

The volume presents an essential selection collected from the essays of Wolfgang Klein. In addition to journal and book articles, many of them published by Mouton, this book features new and unpublished texts by the author. It focuses, among other topics, on information structure, the expression of grammatical categories and the structure of learner varieties.

Abstract

The interaction of action-related language processing with actual movement is an indicator of the functional role of motor cortical involvement in language understanding. This paper describes two experiments using single action verb stimuli. Motor responses were performed with the hand or the foot. To test the double dissociation of language-motor facilitation effects within subjects, Experiments 1 and 2 used a priming procedure where both hand and foot reactions had to be performed in response to different geometrical shapes, which were preceded by action verbs. In Experiment 1, the semantics of the verbs could be ignored whereas Experiment 2 included semantic decisions. Only Experiment 2 revealed a clear double dissociation in reaction times: reactions were facilitated when preceded by verbs describing actions with the matching effector. In Experiment 1, by contrast, there was an interaction between verb-response congruence and a semantic variable related to motor features of the verbs. Thus, the double dissociation paradigm of semantic motor priming was effective, corroborating the role of the motor system in action-related language processing. Importantly, this effect was body part specific.

Abstract

Zwaan et al. mention that young researchers should conduct replications as a
small part of their portfolio. We extend this proposal and suggest that conducting and
reporting replications should become an integral part of PhD projects and be taken into
account in their assessment. We discuss how this would help not only scientific
advancement, but also PhD candidates’ careers.

Abstract

The Dravidian language family consists of about 80 varieties (Hammarström H. 2016 Glottolog 2.7) spoken by 220 million people across southern and central India and surrounding countries (Steever SB. 1998 In The Dravidian languages (ed. SB Steever), pp. 1–39: 1). Neither the geographical origin of the Dravidian language homeland nor its exact dispersal through time are known. The history of these languages is crucial for understanding prehistory in Eurasia, because despite their current restricted range, these languages played a significant role in influencing other language groups including Indo-Aryan (Indo-European) and Munda (Austroasiatic) speakers. Here, we report the results of a Bayesian phylogenetic analysis of cognate-coded lexical data, elicited first hand from native speakers, to investigate the subgrouping of the Dravidian language family, and provide dates for the major points of diversification. Our results indicate that the Dravidian language family is approximately 4500 years old, a finding that corresponds well with earlier linguistic and archaeological studies. The main branches of the Dravidian language family (North, Central, South I, South II) are recovered, although the placement of languages within these main branches diverges from previous classifications. We find considerable uncertainty with regard to the relationships between the main branches.

Abstract

Spatial navigation is a crucial ability for living. Previous animal studies have shown that the S100B gene is causally related to spatial navigation performance in mice. However, the genetic factors influencing human navigation and its neural substrates remain unclear. Here, we provided the first evidence that the S100B gene modulates neural processing of navigationally relevant scenes in humans. First, with a novel protocol, we demonstrated that the spatial pattern of S100B gene expression in postmortem brains was associated with brain activation pattern for spatial navigation in general, and for scene processing in particular. Further, in a large fMRI cohort of healthy adults of Han Chinese (N = 202), we found that S100B gene polymorphisms modulated scene selectivity in the retrosplenial cortex (RSC) and parahippocampal place area. Finally, the serum levels of S100B protein mediated the association between S100B gene polymorphism and scene selectivity in the RSC. Our study takes the first step toward understanding the neurogenetic mechanism of human spatial navigation and suggests a novel approach to discover candidate genes modulating cognitive functions.

Abstract

Spatial navigation is a crucial ability for living. Previous work has revealed multiple distributed brain regions associated with human navigation. However, little is known about how these regions work together as a network (referred to as navigation network) to support flexible navigation. In a novel protocol, we combined neuroimaging meta-analysis, and functional connectivity and behavioral data from the same subjects. Briefly, we first constructed the navigation network for each participant, by combining a large-scale neuroimaging meta-analysis (with the Neurosynth) and resting-state functional magnetic resonance imaging. Then, we investigated multiple topological properties of the navigation networks, including small-worldness, modularity, and highly connected hubs. Finally, we explored the behavioral relevance of these intrinsic properties in a large sample of healthy young adults (N = 190). We found that navigation networks showed small-world and modular organization at global level. More importantly, we found that increased small-worldness and modularity of the navigation network were associated with better navigation ability. Finally, we found that the right retrosplenial complex (RSC) acted as one of the hubs in the navigation network, and that higher betweenness of this region correlated with better navigation ability, suggesting a critical role of the RSC in modulating the navigation network in human brain. Our study takes one of the first steps toward understanding the underlying organization of the navigation network. Moreover, these findings suggest the potential applications of the novel approach to investigating functionally meaningful networks in human brain and their relations to the behavioral impairments in the aging and psychiatric patients.

Abstract

Hemispheric asymmetry is a cardinal feature of human brain organization. Altered brain asymmetry has also been linked to some cognitive and neuropsychiatric disorders. Here the ENIGMA consortium presents the largest ever analysis of cerebral cortical asymmetry and its variability across individuals. Cortical thickness and surface area were assessed in MRI scans of 17,141 healthy individuals from 99 datasets worldwide. Results revealed widespread asymmetries at both hemispheric and regional levels, with a generally thicker cortex but smaller surface area in the left hemisphere relative to the right. Regionally, asymmetries of cortical thickness and/or surface area were found in the inferior frontal gyrus, transverse temporal gyrus, parahippocampal gyrus, and entorhinal cortex. These regions are involved in lateralized functions, including language and visuospatial processing. In addition to population-level asymmetries, variability in brain asymmetry was related to sex, age, and intracranial volume. Interestingly, we did not find significant associations between asymmetries and handedness. Finally, with two independent pedigree datasets (N = 1,443 and 1,113, respectively), we found several asymmetries showing significant, replicable heritability. The structural asymmetries identified, and their variabilities and heritability provide a reference resource for future studies on the genetic basis of brain asymmetry and altered laterality in cognitive, neurological, and psychiatric disorders.

Abstract

Statistical inference plays a critical role in modern scientific research, however, the dominant method for statistical inference in science, null hypothesis significance testing (NHST), is often misunderstood and misused, which leads to unreproducible findings. To address this issue, researchers propose to adopt the Bayes factor as an alternative to NHST. The Bayes factor is a principled Bayesian tool for model selection and hypothesis testing, and can be interpreted as the strength for both the null hypothesis H0 and the alternative hypothesis H1 based on the current data. Compared to NHST, the Bayes factor has the following advantages: it quantifies the evidence that the data provide for both the H0 and the H1, it is not “violently biased” against H0, it allows one to monitor the evidence as the data accumulate, and it does not depend on sampling plans. Importantly, the recently developed open software JASP makes the calculation of Bayes factor accessible for most researchers in psychology, as we demonstrated for the t-test. Given these advantages, adopting the Bayes factor will improve psychological researchers’ statistical inferences. Nevertheless, to make the analysis more reproducible, researchers should keep their data analysis transparent and open.

Abstract

Spatial navigation is a crucial ability for living. Previous studies have shown that males are better at navigation than females, but little is known about the neural basis underlying the sex differences. In this study, we investigated whether cortical scene processing in three well-established scene-selective regions was sexually different, by examining sex differences in scene selectivity and its behavioral relevance to navigation. To do this, we used functional magnetic resonance imaging (fMRI) to scan the parahippocampal place area (PPA), retrosplenial complex (RSC), and occipital place area (OPA) in a large cohort of healthy young adults viewing navigationally relevant scenes (N = 202), and correlated their neural selectivity to scenes with their self-reported navigational ability. Behaviorally, we replicated the previous finding that males were better at navigation than females. Neurally, we found that the scene selectivity in the bilateral PPA, not in the RSC or OPA, was significantly higher in males than females. Such differences could not be explained by confounding factors including brain size and fMRI data quality. Importantly, males, not females, with stronger scene selectivity in the left PPA possessed better navigational ability. This brain-behavior association could not be accounted for by non-navigational abilities (i.e., intelligence and mental rotation ability). Overall, our study provides novel empirical evidence demonstrating sex differences in the brain activity, inviting further studies on sex differences in the neural network for spatial navigation.

Abstract

The leading theories of sentence planning – Hierarchical Incrementality and Linear Incrementality – differ in their assumptions about the coordination of processes that map preverbal information onto language. Previous studies showed that, in native (L1) speakers, this coordination can vary with the ease of executing the message-level and sentence-level processes necessary to plan and produce an utterance. We report the first series of experiments to systematically examine how linguistic experience influences sentence planning in native (L1) speakers (i.e., speakers with life-long experience using the target language) and non-native (L2) speakers (i.e., speakers with less experience using the target language). In all experiments, speakers spontaneously generated one-sentence descriptions of simple events in Dutch (L1) and English (L2). Analyses of eye-movements across early and late time windows (pre- and post-400 ms) compared the extent of early message-level encoding and the onset of linguistic encoding. In Experiment 1, speakers were more likely to engage in extensive message-level encoding and to delay sentence-level encoding when using their L2. Experiments 2–4 selectively facilitated encoding of the preverbal message, encoding of the agent character (i.e., the first content word in active sentences), and encoding of the sentence verb (i.e., the second content word in active sentences) respectively. Experiment 2 showed that there is no delay in the onset of L2 linguistic encoding when speakers are familiar with the events. Experiments 3 and 4 showed that the delay in the onset of L2 linguistic encoding is not due to speakers delaying encoding of the agent, but due to a preference to encode information needed to select a suitable verb early in the formulation process. Overall, speakers prefer to temporally separate message-level from sentence-level encoding and to prioritize encoding of relational information when planning L2 sentences, consistent with Hierarchical Incrementality

Abstract

In the last decade, the involvement of neural oscillatory mechanisms in speech comprehension has been increasingly investigated. Current evidence suggests that low-frequency and high-frequency neural entrainment to the acoustic dynamics of speech are linked to its analysis. One crucial question is whether acoustical processing primarily modulates neural entrainment, or whether entrainment instead reflects linguistic processing. Here, we review studies investigating the effect of linguistic manipulations on neural oscillatory activity. In light of the current findings, we argue that theta (3–8 Hz) entrainment may primarily reflect the analysis of the acoustic features of speech. In contrast, recent evidence suggests that delta (1–3 Hz) and high-frequency activity (>40 Hz) are reliable indicators of perceived linguistic representations. The interdependence between low-frequency and high-frequency neural oscillations, as well as their causal role on speech comprehension, is further discussed with regard to neurophysiological models of speech processing

Abstract

Low-frequency neural entrainment to rhythmic input
has been hypothesized as a canonical mechanism
that shapes sensory perception in time. Neural
entrainment is deemed particularly relevant for
speech analysis, as it would contribute to the extraction
of discrete linguistic elements from continuous
acoustic signals. However, its causal influence in
speech perception has been difficult to establish.
Here, we provide evidence that oscillations build temporal
predictions about the duration of speech tokens
that affect perception. Using magnetoencephalography
(MEG), we studied neural dynamics during
listening to sentences that changed in speech rate.
Weobserved neural entrainment to preceding speech
rhythms persisting for several cycles after the change
in rate. The sustained entrainment was associated
with changes in the perceived duration of the last
word’s vowel, resulting in the perception of words
with different meanings. These findings support oscillatory
models of speech processing, suggesting that
neural oscillations actively shape speech perception.

Abstract

Research questions:

Are emergent bilinguals sensitive to register variation in their use of communication strategies? What strategies do LX speakers, in casu Spanish speakers of English, use as a function of situational context? What role do individual differences play?
Methodology:

This within-speaker study compares Spanish second-language English speakers’ communication strategy use in an informal, peer-to-peer conversation and a formal interview.
Data and analysis:

The 15 hours of informal and 9.5 hours of formal speech from the Nijmegen Corpus of Spanish English were coded for 19 different communication strategies.
Findings/conclusions:

Overall, speakers prefer self-reliant strategies, which allow them to continue communication without their interlocutor’s help. Of the self-reliant strategies, least effort strategies such as code-switching are used more often in informal speech, whereas relatively more effortful strategies (e.g. reformulations) are used more in informal speech, when the need to be unambiguously understood is felt as more important. Individual differences played a role: some speakers were more affected by a change in formality than others.
Originality:

Sensitivity to register variation has not yet been studied within communicative strategy use.
Implications:

General principles of communication govern speakers’ strategy selection, notably the protection of positive face and the least effort and cooperative principles.

Abstract

English serves as a lingua franca in situations with varying degrees of
formality. How formality affects non-native speech has rarely been studied. We
investigated register variation by Spanish users of English by comparing formal
and informal speech from the Nijmegen Corpus of Spanish English that we
created. This corpus comprises speech from thirty-four Spanish speakers of
English in interaction with Dutch confederates in two speech situations.
Formality affected the amount of laughter and overlapping speech and the
number of Spanish words. Moreover, formal speech had a more informational
character than informal speech. We discuss how our findings relate to register
variation in Spanish

Abstract

Background

Left-right asymmetry is a fundamental organizing feature of the human brain, and neuro-psychiatric disorders such as schizophrenia sometimes involve alterations of brain asymmetry. As early as 8 weeks post conception, the majority of human fetuses move their right arms more than their left arms, but because nerve fibre tracts are still descending from the forebrain at this stage, spinal-muscular asymmetries are likely to play an important developmental role.
Methods

We used RNA sequencing to measure gene expression levels in the left and right spinal cords, and left and right hindbrains, of 18 post-mortem human embryos aged 4-8 weeks post conception. Genes showing embryonic lateralization were tested for an enrichment of signals in genome-wide association data for schizophrenia.
Results

The left side of the embryonic spinal cord was found to mature faster than the right side. Both sides transitioned from transcriptional profiles associated with cell division and proliferation at earlier stages, to neuronal differentiation and function at later stages, but the two sides were not in synchrony (p = 2.2 E-161). The hindbrain showed a left-right mirrored pattern compared to the spinal cord, consistent with the well-known crossing over of function between these two structures. Genes that showed lateralization in the embryonic spinal cord were enriched for association signals with schizophrenia (p = 4.3 E-05).
Conclusions
These are the earliest-stage left-right differences of human neural development ever reported. Disruption of the lateralised developmental programme may play a role in the genetic susceptibility to schizophrenia.

Abstract

Importance Knowing the range of symptoms seen in patients with a missense or loss-of-function variant in KCNB1 and how these symptoms correlate with the type of variant will help clinicians with diagnosis and prognosis when treating new patients. Objectives To investigate the clinical spectrum associated with KCNB1 variants and the genotype-phenotype correlations. Design, Setting, and Participants This study summarized the clinical and genetic information of patients with a presumed pathogenic variant in KCNB1.Patients were identified in research projects or during clinical testing. Information on patients from previously published articles was collected and authors contacted if feasible. All patients were seen at a clinic at one of the participating institutes because of presumed genetic disorder. They were tested in a clinical setting or included in a research project. Main Outcomes and Measures The genetic variant and its inheritance and information on the patient's symptoms and characteristics in a predefined format. All variants were identified with massive parallel sequencing and confirmed with Sanger sequencing in the patient. Absence of the variant in the parents could be confirmed with Sanger sequencing in all families except one. Results Of 26 patients (10 female, 15 male, 1 unknown; mean age at inclusion, 9.8 years; age range, 2-32 years) with developmental delay, 20 (77%) carried a missense variant in the ion channel domain of KCNB1, with a concentration of variants in region S5 to S6. Three variants that led to premature stops were located in the C-terminal and 3 in the ion channel domain. Twenty-one of 25 patients (84%) had seizures, with 9 patients (36%) starting with epileptic spasms between 3 and 18 months of age. All patients had developmental delay, with 17 (65%) experiencing severe developmental delay; 14 (82%) with severe delay had behavioral problems. The developmental delay was milder in 4 of 6 patients with stop variants and in a patient with a variant in the S2 transmembrane element rather than the S4 to S6 region. Conclusions and Relevance De novo KCNB1 missense variants in the ion channel domain and loss-of-function variants in this domain and the C-terminal likely cause neurodevelopmental disorders with or without seizures. Patients with presumed pathogenic variants in KCNB1 have a variable phenotype. However, the type and position of the variants in the protein are (imperfectly) correlated with the severity of the disorder.

Abstract

Left-right laterality is an important aspect of human –and in fact all vertebrate– brain organization for which the genetic basis is poorly understood. Using RNA sequencing data we contrasted gene expression in left- and right-sided samples from several structures of the anterior central nervous systems of post mortem human embryos and foetuses. While few individual genes stood out as significantly lateralized, most structures showed evidence of laterality of their overall transcriptomic profiles. These left-right differences showed overlap with age-dependent changes in expression, indicating lateralized maturation rates, but not consistently in left-right orientation over all structures. Brain asymmetry may therefore originate in multiple locations, or if there is a single origin, it is earlier than 5 weeks post conception, with structure-specific lateralized processes already underway by this age. This pattern is broadly consistent with the weak correlations reported between various aspects of adult brain laterality, such as language dominance and handedness.

Abstract

Left-right asymmetry is subtle but pervasive in the human central nervous system. This asymmetry is initiated early during development, but its mechanisms are poorly known. Forebrains and midbrains were dissected from six human embryos at Carnegie stages 15 or 16, one of which was female. The structures were divided into left and right sides, and RNA was isolated. RNA was sequenced with 100 base-pair paired ends using Illumina Hiseq 4000. After quality control, five paired brain sides were available for midbrain and forebrain. A paired analysis between left- and right sides of a given brain structure across the embryos identified left-right differences. The dataset, consisting of Fastq files and a read count table, can be further used to study early development of the human brain

Abstract

The intercalated cells (ITCs) of the amygdala have been shown to be critical regulatory components of amygdalar circuits, which control appropriate fear responses. Despite this, the molecular processes guiding ITC development remain poorly understood. Here we establish the zinc finger transcription factor Tshz1 as a marker of ITCs during their migration from the dorsal lateral ganglionic eminence through maturity. Using germline and conditional knock-out (cKO) mouse models, we show that Tshz1 is required for the proper migration and differentiation of ITCs. In the absence of Tshz1, migrating ITC precursors fail to settle in their stereotypical locations encapsulating the lateral amygdala and BLA. Furthermore, they display reductions in the ITC marker Foxp2 and ectopic persistence of the dorsal lateral ganglionic eminence marker Sp8. Tshz1 mutant ITCs show increased cell death at postnatal time points, leading to a dramatic reduction by 3 weeks of age. In line with this, Foxp2-null mutants also show a loss of ITCs at postnatal time points, suggesting that Foxp2 may function downstream of Tshz1 in the maintenance of ITCs. Behavioral analysis of male Tshz1 cKOs revealed defects in fear extinction as well as an increase in floating during the forced swim test, indicative of a depression-like phenotype. Moreover, Tshz1 cKOs display significantly impaired social interaction (i.e., increased passivity) regardless of partner genetics. Together, these results suggest that Tshz1 plays a critical role in the development of ITCs and that fear, depression-like and social behavioral deficits arise in their absence. SIGNIFICANCE STATEMENT We show here that the zinc finger transcription factor Tshz1 is expressed during development of the intercalated cells (ITCs) within the mouse amygdala. These neurons have previously been shown to play a crucial role in fear extinction. Tshz1 mouse mutants exhibit severely reduced numbers of ITCs as a result of abnormal migration, differentiation, and survival of these neurons. Furthermore, the loss of ITCs in mouse Tshz1 mutants correlates well with defects in fear extinction as well as the appearance of depression-like and abnormal social interaction behaviors reminiscent of depressive disorders observed in human patients with distal 18q deletions, including the Tshz1 locus.

Abstract

In this exploratory study we test the hypothesis that the retrieval from memory of proper noun Agents (PNAs) under processing pressure causes a greater proportion of such semantic arguments to be placed to the right of the initial position in a clause than would be the case if such retrieval from memory were not necessary. This effect is manifest in sports commentary. Processing pressure on sports commentators is modulated by the speed at which the sport is played and reported. Non-initial placement is also facilitated by formulae which have slots in non-initial position. It follows that the non-initial placement of PNAs is not always semantically or pragmatically motivated. This finding therefore runs counter to a strong form of the functionalist hypothesis that syntactic choices available in the systemic structure of the syntax of a language offer solely semantic or pragmatic choices. It is an open question in a weak functionalist account of language and language use how processing and communicative functions interact in general.

Abstract

Many natural auditory signals, including music and language, change periodically. The effect of such auditory rhythms on the brain is unclear however. One widely held view, dynamic attending theory, proposes that the attentional system entrains to the rhythm and increases attention at moments of rhythmic salience. In support, 2 experiments reported here show reduced response times to visual letter strings shown at auditory rhythm peaks, compared with rhythm troughs. However, we argue that an account invoking the entrainment of general attention should further predict rhythm entrainment to also influence memory for visual stimuli. In 2 pseudoword memory experiments we find evidence against this prediction. Whether a pseudoword is shown during an auditory rhythm peak or not is irrelevant for its later recognition memory in silence. Other attention manipulations, dividing attention and focusing attention, did result in a memory effect. This raises doubts about the suggested attentional nature of rhythm entrainment. We interpret our findings as support for auditory rhythm perception being based on auditory-motor entrainment, not general attention entrainment.

Abstract

In response to recommendations to redefine statistical significance to P ≤ 0.005, we propose that researchers should transparently report and justify all choices they make when designing a study, including the alpha level.

Abstract

The current electroencephalography study investigated the relationship between the motor and (language) comprehension systems by simultaneously measuring mu and N400 effects. Specifically, we examined whether the pattern of motor activation elicited by verbs depends on the larger sentential context. A robust N400 congruence effect confirmed the contextual manipulation of action plausibility, a form of semantic congruency. Importantly, this study showed that: (1) Action verbs elicited more mu power decrease than non-action verbs when sentences described plausible actions. Action verbs thus elicited more motor activation than non-action verbs. (2) In contrast, when sentences described implausible actions, mu activity was present but the difference between the verb types was not observed. The increased processing associated with a larger N400 thus coincided with mu activity in sentences describing implausible actions. Altogether, context-dependent motor activation appears to play a functional role in deriving context-sensitive meaning

Abstract

Neural oscillations may be instrumental for the tracking and segmentation of continuous speech. Earlier work has suggested that delta, theta and gamma oscillations entrain to the speech rhythm. We used magnetoencephalography and a large sample of 102 participants to investigate oscillatory entrainment to speech, and observed robust entrainment of delta and theta activity, and weak group-level gamma entrainment. We show that the peak frequency and the hemispheric lateralisation of the entrainment are subject to considerable individual variability. The first finding may support the involvement of intrinsic oscillations in entrainment, and the second finding suggests that there is no systematic default right-hemispheric bias for processing acoustic signals on a slow time scale. Although low frequency entrainment to speech is a robust phenomenon, the characteristics of entrainment vary across individuals, and this variation is important for understanding the underlying neural mechanisms of entrainment, as well as its functional significance.

Abstract

Sensory systems experience a trade-off between maximizing the
detail and amount of sampled information. Thistrade-off is particularly
pronounced in sensorysystemsthat are highlyspecialised fora single
task and thus experience limitations in other tasks. We hypothesised
that combining sensory input from multiple streams of information
may resolve this trade-off and improve detection and sensing
reliability. Specifically, we predicted that perceptive limitations
experienced by animals reliant on specialised active echolocation
can be compensated for by the phylogenetically older and less
specialised process of passive hearing. We tested this hypothesis in
greater horseshoe bats, which possess morphological and neural
specialisations allowing them to identify fluttering prey in dense
vegetation using echolocation only. At the same time, their
echolocation system is both spatially and temporally severely
limited. Here, we show that greater horseshoe bats employ passive
hearing to initially detect and localise prey-generated and other
environmental sounds, and then raise vocalisation level and
concentrate the scanning movements of their sonar beam on the
sound source for further investigation with echolocation. These
specialised echolocators thus supplement echo-acoustic information
with environmental acoustic cues, enlarging perceived space beyond
their biosonar range. Contrary to our predictions, we did not find
consistent preferences for prey-related acoustic stimuli, indicating the
use of passive acoustic cues also for detection of non-prey objects.
Our findings suggest that even specialised echolocators exploit a
wide range of environmental information, and that phylogenetically
older sensory systems can support the evolution of sensory
specialisations by compensating for their limitations.