Publications

Abstract

This article presents a psycholinguistically inspired approach to the syntax of clause-level coordination and coordinate ellipsis. It departs from the assumption that coordinations are structurally similar to so-called appropriateness repairs — an important type of self-repairs in spontaneous speech. Coordinate structures and appropriateness repairs can both be viewed as “update” constructions. Updating is defined as a special sentence production mode that efficiently revises or augments existing sentential structure in response to modifications in the speaker's communicative intention. This perspective is shown to offer an empirically satisfactory and theoretically parsimonious account of two prominent types of coordinate ellipsis, in particular “forward conjunction reduction” (FCR) and “gapping” (including “long-distance gapping” and “subgapping”). They are analyzed as different manifestations of “incremental updating” — efficient updating of only part of the existing sentential structure. Based on empirical data from Dutch and German, novel treatments are proposed for both types of clausal coordinate ellipsis. The coordination-as-updating perspective appears to explain some general properties of coordinate structure: the existence of the well-known “coordinate structure constraint”, and the attractiveness of three-dimensional representations of coordination. Moreover, two other forms of coordinate ellipsis — SGF (“subject gap in finite clauses with fronted verb”), and “backward conjunction reduction” (BCR) (also known as “right node raising” or RNR) — are shown to be incompatible with the notion of incremental updating. Alternative theoretical interpretations of these phenomena are proposed. The four types of clausal coordinate ellipsis — SGF, gapping, FCR and BCR — are argued to originate in four different stages of sentence production: Intending (i.e., preparing the communicative intention), conceptualization, grammatical encoding, and phonological encoding, respectively.

Abstract

This paper presents a theory of the syntactic aspects of human sentence production. An important characteristic of unprepared speech is that overt pronunciation of a sentence can be initiated before the speaker has completely worked out the meaning content he or she is going to express in that sentence. Apparently, the speaker is able to build up a syntactically coherent utterance out of a series of syntactic fragments each rendering a new part of the meaning content. This incremental, left-to-right mode of sentence production is the central capability of the proposed Incremental Procedural Grammar (IPG). Certain other properties of spontaneous speech, as derivable from speech errors, hesitations, self-repairs, and language pathology, are accounted for as well. The psychological plausibility thus gained by the grammar appears compatible with a satisfactory level of linguistic plausibility in that sentences receive structural descriptions which are in line with current theories of grammar. More importantly, an explanation for the existence of configurational conditions on transformations and other linguistics rules is proposed. The basic design feature of IPG which gives rise to these psychologically and linguistically desirable properties, is the “Procedures + Stack” concept. Sentences are built not by a central constructing agency which overlooks the whole process but by a team of syntactic procedures (modules) which work-in parallel-on small parts of the sentence, have only a limited overview, and whose sole communication channel is a stock. IPG covers object complement constructions, interrogatives, and word order in main and subordinate clauses. It handles unbounded dependencies, cross-serial dependencies and coordination phenomena such as gapping and conjunction reduction. It is also capable of generating self-repairs and elliptical answers to questions. IPG has been implemented as an incremental Dutch sentence generator written in LISP.

Abstract

Human speakers often produce sentences incrementally. They can start speaking having in mind only a fragmentary idea of what they want to say, and while saying this they refine the contents underlying subsequent parts of the utterance. This capability imposes a number of constraints on the design of a syntactic processor. This paper explores these constraints and evaluates some recent computational sentence generators from the perspective of incremental production.

Abstract

[Seven one-page psycholinguistic sketches]

Abstract

The Max Planck Institute for Psycholinguistics in Nijmegen, The Netherlands, is creating a state-of-the-art web environment for the ISO TC 37 (terminology and other language and content resources) metadata registry. This Data Category Registry (DCR) is called ISOcat and encompasses data categories for a broad range of language resources. Under the governance of the DCR Board, ISOcat provides an open work space for creating data category specifications, defining Data Category Selections (DCSs) (domain-specific groups of data categories), and standardising selected data categories and DCSs. Designers visualise future interactivity among the DCR, reference registries and ontological knowledge spaces

Abstract

Spatial terms in the world’s languages appear to reflect both universal conceptual tendencies and linguistic convention. A similarly mixed picture in the case of color naming has been accounted for in terms of near-optimal partitions of color space. Here, we demonstrate that this account generalizes to spatial terms. We show that the spatial terms of 9 diverse languages near-optimally partition a similarity space of spatial meanings, just as color terms near-optimally partition color space. This account accommodates both universal tendencies and cross-language differences in spatial category extension, and identifies general structuring principles that appear to operate across different semantic domains.

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

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

In the first, critical part of this study, a small sample of simple German sentences with their empirically determined pitch contours is used to demonstrate the incorrectness of numerous currently hold views of German sentence intonation. In the second, more constructive part, several interrogative sentence types are analysed and an attempt is made to show that intonation, besides other functions, indicates the permantently changing 'thematic score' in on-going discourse as well as certain validity claims.

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

Nobel's legacy, or the metamorphosis of what is idealistic Ever since the first Nobel prize in literature was awarded to Prudhomme in 1901, the decisions of the Swedish Academy have been subject to criticism. What is surprising in the changing decision policy as well as in its criticism is the fact that Alfred Nobel's original intentions are hardly ever taken into account: the Nobel prize is a philanthropic prize, it is not meant to select and honour the most eminent literary work but the work with maximal benefit to human beings. What is even more surprising is the fact that no one seems to care that the donator's Last Will is regularly broken.

Abstract

In this article, we discuss the implications of the fact that adult second language learners (outside the classroom) universally develop a well-structured, efficient and simple form of language–the Basic Variety (BV). Three questions are asked as to (1) the structural properties of the BV, (2) the status of these properties and (3) why some structural properties of ‘fully fledged’ languages are more complex. First, we characterize the BV in four respects: its lexical repertoire, the principles according to which utterances are structured, and temporality and spatiality expressed. The organizational principles proposed are small in number, and interact. We analyse this interaction, describing how the BV is put to use in various complex verbal tasks, in order to establish both what its communicative potentialities are, and also those discourse contexts where the constraints come into conflict and where the variety breaks down. This latter phenomenon provides a partial answer to the third question,concerning the relative complexity of ‘fully fledged’ languages–they have devices to deal with such cases. As for the second question, it is argued firstly that the empirically established continuity of the adult acquisition process precludes any assignment of the BV to a mode of linguistic expression (e.g., ‘protolanguage’) distinct from that of ‘fully fledged’ languages and, moreover, that the organizational constraints of the BV belong to the core attributes of the human language capacity, whereas a number of complexifications not attested in the BV are less central properties of this capacity. Finally, it is shown that the notion of feature strength, as used in recent versions of Generative Grammar, allows a straightforward characterization of the BV as a special case of an I-language, in the sense of this theory. Under this perspective, the acquisition of an Ilanguage beyond the BV can essentially be described as a change in feature strength.

Abstract

Starting with Ross (1970), various proposals have been made to classify elliptical coordinations and to characterize different languages according to the types of ellipses which they admit. Sanders (1977) discusses four of these proposals, shows that they are inadequate on various grounds and proposes a fifth typology whose central claim is 'evidently correct', as he states (p. 258). In the following, I shall briefly outline this typology and then show that it is inadequate, too. Since there is only one language 1 know — German — I will take all my examples from this language. Moreover, all examples will be straightforward and easy to be judged.

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 Max Planck Institute for Psycholinguistics (MPI) manages an archive of linguistic research data with a current size of almost 20 Terabytes. Apart from in-house researchers other projects also store their data in the archive, most notably the Documentation of Endangered Languages (DoBeS) projects. The archive is available online and can be accessed by anybody with Internet access. To be able to manage this large amount of data the MPI's technical group has developed a software suite called Language Archiving Technology (LAT) that on the one hand helps researchers and archive managers to manage the data and on the other hand helps users in enriching their primary data with additional layers. All the MPI software is Java-based and developed according to open source principles (GNU, 2007). All three major operating systems (Windows, Linux, MacOS) are supported and the software works similarly on all of them. As the archive is online, many of the tools, especially the ones for accessing the data, are browser based. Some of these browser-based tools make use of Adobe Flex to create nice-looking GUIs. The LAT suite is a complete set of management and enrichment tools, and given the interaction between the tools the result is a complete LAT software domain. Over the last 10 years, this domain has proven its functionality and use, and is being deployed to servers in other institutions. This deployment is an important step in getting the archived resources back to the members of the speech communities whose languages are documented. In the paper we give an overview of the tools of the LAT suite and we describe their functionality and role in the integrated process of archiving, management and enrichment of linguistic data.

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

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

To compare abstract structural and lexicalist accounts of syntactic processes in sentence formulation, we examined the effectiveness of nonidiomatic and idiomatic phrasal verbs in inducing structural generalizations. Three experiments made use of a syntactic priming paradigm in which participants recalled sentences they had read in rapid serial visual presentation. Prime and target sentences contained phrasal verbs with particles directly following the verb (pull off a sweatshirt) or following the direct object (pull a sweatshirt off). Idiomatic primes used verbs whose figurative meaning cannot be straightforwardly derived from the literal meaning of the main verb (e.g., pull off a robbery) and are commonly treated as stored lexical units. Particle placement in sentences was primed by both nonidiomatic and idiomatic verbs. Experiment 1 showed that the syntax of idiomatic and nonidiomatic phrasal verbs is amenable to priming, and Experiments 2 and 3 compared the priming patterns created by idiomatic and nonidiomatic primes. Despite differences in idiomaticity and structural flexibility, both types of phrasal verbs induced structural generalizations and differed little in their ability to do so. The findings are interpreted in terms of the role of abstract structural processes in language production.

Abstract

Source monitoring can be influenced by information that is external to the study context, such as beliefs and general knowledge (Johnson, Hashtroudi, & Lindsay, 1993). We investigated the extent to which metamnemonic judgments predict memory for items and sources when schematic information about the sources is or is not provided at encoding. Participants made judgments of learning (JOLs) to statements presented by two speakers and were informed of the occupation of each speaker either before or after the encoding session. Replicating earlier work, prior knowledge decreased participants' tendency to erroneously attribute statements to schematically consistent but episodically incorrect speakers. The origin of this effect can be understood by examining the relationship between JOLs and performance: JOLs were equally predictive of item and source memory in the absence of prior knowledge, but were exclusively predictive of source memory when participants knew of the relationship between speakers and statements during study. Background knowledge determines the information that people solicit in service of metamnemonic judgments, suggesting that these judgments reflect control processes during encoding that reduce schematic errors.

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

Recognizing word boundaries in continuous speech requires detailed knowledge of the native language. In the first year of life, infants acquire considerable word segmentation abilities. Infants at this early stage in word segmentation rely to a large extent on the metrical pattern of their native language, at least in stress-based languages. In Dutch and English (both languages with a preferred trochaic stress pattern), segmentation of strong-weak words develops rapidly between 7 and 10 months of age. Nevertheless, trochaic languages contain not only strong-weak words but also words with a weak-strong stress pattern. In this article, we present electrophysiological evidence of the beginnings of weak-strong word segmentation in Dutch 10-month-olds. At this age, the ability to combine different cues for efficient word segmentation does not yet seem to be completely developed. We provide evidence that Dutch infants still largely rely on strong syllables, even for the segmentation of weak-strong words.

Abstract

The paper investigates the use of manner verbs (e.g. marcher 'to walk', courir 'to run') with so-called locative prepositions (e.g. dans 'in', sous 'under') in the descriptions of motion in French, as in Il a couru dans le bureau 'He ran in (to) the office', to explore the type of events such constructions express and the factors that influence their interpretation. Based on an extensive corpus survey, the study shows that, contrary to the general claim according to which such constructions express typically motion in some location, they are also frequently used to express change of location. The study discusses the interplay of various factors that contribute to the interpretation of these constructions, including semantic, aspectual and pragmatic factors.

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

Words can be distinguished by segmental differences or by suprasegmental differences or both. Studies from English suggest that suprasegmentals play little role in human spoken-word recognition; English stress, however, is nearly always unambiguously coded in segmental structure (vowel quality); this relationship is less close in Dutch. The present study directly compared the effects of segmental and suprasegmental mispronunciation on word recognition in Dutch. There was a strong effect of suprasegmental mispronunciation, suggesting that Dutch listeners do exploit suprasegmental information in word recognition. Previous findings indicating the effects of mis-stressing for Dutch differ with stress position were replicated only when segmental change was involved, suggesting that this is an effect of segmental rather than suprasegmental processing.

Abstract

Little is known about the genetic contribution to individual differences in neural networks subserving cognition function. In this functional magnetic resonance imaging (fMRI) twin study, we found a significant genetic influence on brain activation in neural networks supporting digit working memory tasks. Participants activating frontal-parietal networks responded faster than individuals relying more on language-related brain networks.There were genetic influences on brain activation in language-relevant brain circuits that were atypical for numerical working memory tasks as such. This suggests that differences in cognition might be related to brain activation patterns that differ qualitatively among individuals.

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

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

Heterozygous mutations of the human FOXP2 gene cause a developmental disorder involving impaired learning and production of fluent spoken language. Previous investigations of its aetiology have focused on disturbed function of neural circuits involved in motor control. However, Foxp2 expression has been found in the cochlea and auditory brain centers and deficits in auditory processing could contribute to difficulties in speech learning and production. Here, we recorded auditory brainstem responses (ABR) to assess two heterozygous mouse models carrying distinct Foxp2 point mutations matching those found in humans with FOXP2-related speech/language impairment. Mice which carry a Foxp2-S321X nonsense mutation, yielding reduced dosage of Foxp2 protein, did not show systematic ABR differences from wildtype littermates. Given that speech/language disorders are observed in heterozygous humans with similar nonsense mutations (FOXP2-R328X), our findings suggest that auditory processing deficits up to the midbrain level are not causative for FOXP2-related language impairments. Interestingly, however, mice harboring a Foxp2-R552H missense mutation displayed systematic alterations in ABR waves with longer latencies (significant for waves I, III, IV) and smaller amplitudes (significant for waves I, IV) suggesting that either the synchrony of synaptic transmission in the cochlea and in auditory brainstem centers is affected, or fewer auditory nerve fibers and fewer neurons in auditory brainstem centers are activated compared to wildtypes. Therefore, the R552H mutation uncovers possible roles for Foxp2 in the development and/or function of the auditory system. Since ABR audiometry is easily accessible in humans, our data call for systematic testing of auditory functions in humans with FOXP2 mutations.