Publications

Abstract

In this study, we use the association between various measures of the morphological family and decision latencies to reveal the way in which the components of Dutch and English compounds are processed. The results show that for constituents of concatenated compounds in both languages, a position-related token count of the morphological family plays a role, whereas English open compounds show an effect of a type count, similar to the effect of family size for simplex words. When Dutch compounds are written with an artificial space, they reveal no effect of type count, which shows that the differential effect for the English open compounds is not superficial. The final experiment provides converging evidence for the lexical consequences of the space in English compounds. Decision latencies for English simplex words are better predicted from counts of the morphological family that include concatenated and hyphenated but not open family members.

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

Day-to-day interpretations of “space” are enmeshed in specific cultural and linguistic practices. For example, many cultures have an association between vertical height and social standing; more powerful people may be placed literally higher than others at social gatherings, and be spoken of as having higher status. This questionnaire is a guide for exploring relationships between space, language, and social structure. The goal is to better understand how space is organised in the focus community, and to investigate the extent to which space is used as a model for reproducing social forms.

Abstract

In this paper we present a definition of Performance Grammar (PG), a psycholinguistically motivated syntax formalism, in declarative terms. PG aims not only at describing and explaining intuitive judgments and other data concerning the well–formedness of sentences of a language, but also at contributing to accounts of syntactic processing phenomena observable in language comprehension and language production. We highlight two general properties of human sentence generation, incrementality and late linearization,which make special demands on the design of grammar formalisms claiming psychological plausibility. In order to meet these demands, PG generates syntactic structures in a two-stage process. In the first and most important ‘hierarchical’ stage, unordered hierarchical structures (‘mobiles’) are assembled out of lexical building blocks. The key operation at work here is typed feature unification, which also delimits the positional options of the syntactic constituents in terms of so-called topological features. The second, much simpler stage takes care of arranging the branches of the mobile from left to right by ‘reading–out’ one positional option of every constituent. In this paper we concentrate on the structure assembly formalism in PG’s hierarchical component. We provide a declarative definition couched in an HPSG–style notation based on typed feature unification. Our emphasis throughout is on linear order constraints.

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

When you compare the behavior of two different age groups which are trying to master the same sensori-motor or cognitive skill, you are likely to discover varying learning routes: different stages, different intervals between stages, or even different orderings of stages. Such heterogeneous learning trajectories may be caused by at least six different types of factors: (1) Initial state: the kinds and levels of skills the learners have available at the onset of the learning episode. (2) Learning mechanisms: rule-based, inductive, connectionist, parameter setting, and so on. (3) Input and feedback characteristics: learning stimuli, information about success and failure. (4) Information processing mechanisms: capacity limitations, attentional biases, response preferences. (5) Energetic variables: motivation, emotional reactions. (6) Final state: the fine-structure of kinds and levels of subskills at the end of the learning episode. This applies to language acquisition as well. First and second language learners probably differ on all six factors. Nevertheless, the debate between advocates and opponents of the Fundamental Difference Hypothesis concerning L1 and L2 acquisition have looked almost exclusively at the first two factors. Those who believe that L1 learners have access to Universal Grammar whereas L2 learners rely on language processing strategies, postulate different learning mechanisms (UG parameter setting in L1, more general inductive strategies in L2 learning). Pienemann opposes this view and, based on his Processability Theory, argues that L1 and L2 learners start out from different initial states: they come to the grammar learning task with different structural hypotheses (SOV versus SVO as basic word order of German).

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

Comments on a study by Frazier and others on Dutch-language lexical processing. Claims that the control condition in the experiment was inadequate and that an assumption made by Frazier about closed class verbal items is inaccurate, and proposes an alternative account of a subset of the data from the experiment

Abstract

Raises objections to L. Frazier et al's (see record 1994-32229-001) report of an experimental study intended to test Schreuder's (1990) Morphological Integration (MI) model concerning the processing of separable and inseparable verbs and shows that the logic of the experiment is flawed. The problem is rooted in the notion of a separable complex verb. The conclusion is drawn that Frazier et al's experimental data cannot be taken as evidence for the theoretical propositions they develop about the MI model.

Abstract

Children must possess some ability to process input in a meaningful manner to acquire language. The present study reports on data from an experiment investigating 3- to 5-year-old English-speaking children's understanding of restrictive relative clauses manipulated for embeddedness and focus. The results of the study showed that English-speaking children acquire right-branching before center-embedded structures. Comparisons made with data from Portuguese-speaking children suggest general-cognitive and language-specific constraints on development, and with respect to English, a “clause expansion” approach to processing in development

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

This task investigates the expression of “enter” and “exit” events, and is a supplement to the Motion Elicitation task (https://doi.org/10.17617/2.3003391). Consultants are asked to describe a series of animated clips where a man moves into or out of a house. The clips focus on contrasts to do with perspective (e.g., whether the man appears to move away or towards the viewer) and transitional movement (e.g., whether the man walks or “teleports” into his new location).

Abstract

Do our hands 'speak the same language' across cultures? Gesture is the silent partner of spoken languages in face-to-face interaction, but we still have a lot to learn about gesture practices in different speech communities. The primary purpose of this task is to collect data in naturalistic settings that can be used to investigate the linguistic and cultural relativity of gesture performance, especially spatially indicative gestures. It involves video-recording pairs of speakers in both free conversation and more structured communication tasks (e.g., describing film plots).

Please note: the stimuli mentioned in this entry are available elsewhere: 'The Pear Story', a short film made at the University of California at Berkeley; "Frog, where are you?" from the original Mayer (1969) book, as published in the Appendix of Berman & Slobin (1994).

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

It is generally assumed that verbs have an ‘argument structure’, which imposes various constraints on the noun phrases that can or must go with the verb, and an ‘event structure’, which characterises the particular temporal characteristics of the ‘event’ which the verb relates to: this event may be a state, a process, an activity, an ‘event in the narrow sense’, and others. In this paper, it is argued that that argument structure and event structure should be brought together. The lexical content of a verb assigns descriptive properties to one or more arguments at one or more times, hence verbs have an ‘argument time-structure’ (AT-structure). Numerous morphological and syntactical operations, such as participle formation or complex verb constructions, modify this AT-structure. This is illustrated with German recipient constructions such as ein Buch geschenkt bekommen or das Fenster geöffnet kriegen.

Abstract

The German Perfekt has two quite different temporal readings, as illustrated by the two possible continuations of the sentence Peter hat gearbeitet in i, ii, respectively: (i) Peter hat gearbeitet und ist müde. Peter has worked and is tired. (ii) Peter hat gearbeitet und wollte nicht gestört werden. Peter has worked and wanted not to be disturbed. The first reading essentially corresponds to the English present perfect; the second can take a temporal adverbial with past time reference ('yesterday at five', 'when the phone rang', and so on), and an English translation would require a past tense ('Peter worked/was working'). This article shows that the Perfekt has a uniform temporal meaning that results systematically from the interaction of its three components-finiteness marking, auxiliary and past participle-and that the two readings are the consequence of a structural ambiguity. This analysis also predicts the properties of other participle constructions, in particular the passive in German.

Abstract

Chinese has a number of particles such as le, guo, zai and zhe that add a particular aspectual value to the verb to which they are attached. There have been many characterisations of this value in the literature. In this paper, we review several existing influential accounts of these particles, including those in Li and Thompson (1981), Smith (1991), and Mangione and Li (1993). We argue that all these characterisations are intuitively plausible, but none of them is precise.We propose that these particles serve to mark which part of the sentence''s descriptive content is asserted, and that their aspectual value is a consequence of this function. We provide a simple and precise definition of the meanings of le, guo, zai and zhe in terms of the relationship between topic time and time of situation, and show the consequences of their interaction with different verb expressions within thisnew framework of interpretation.

Abstract

During the last 25 years, second language acquisition (SLA) research hasmade considerable progress, but is still far from proving a solid basis for foreign language teaching, or from a general theory of SLA. In addition, its status within the linguistic disciplines is still very low. I argue this has not much to do with low empirical or theoretical standards in the field—in this regard, SLA research is fully competitive—but with a particular perspective on the acquisition process: SLA researches learners' utterances as deviations from a certain target, instead of genuine manifestations of underlying language capacity; it analyses them in terms of what they are not rather than what they are. For some purposes such a "target deviation perspective" makes sense, but it will not help SLA researchers to substantially and independently contribute to a deeper understanding of the structure and function of the human language faculty. Therefore, these findings will remain of limited interest to other scientists until SLA researchers consider learner varieties a normal, in fact typical, manifestation of this unique human capacity.

Abstract

Objectives: To investigate the time resolution of different methods for the computation of event-related desynchronization/synchronization (ERD/ERS), including one based on Hilbert transform. Methods: In order to better understand the time resolution of ERD/ERS, which is a function of factors such as the exact computation method, the frequency under study, the number of trials, and the sampling frequency, we simulated sudden changes in oscillation amplitude as well as very short and closely spaced events. Results: Hilbert-based ERD yields very similar results to ERD integrated over predefined time intervals (block ERD), if the block length is half the period length of the studied frequency. ERD predicts the onset of a change in oscillation amplitude with an error margin of only 10–30 ms. On the other hand, the time the ERD response needs to climb to its full height after a sudden change in oscillation amplitude is quite long, i.e. between 200 and 500 ms. With respect to sensitivity to short oscillatory events, the ratio between sampling frequency and electroencephalographic frequency band plays a major role. Conclusions: (1) The optimal time interval for the computation of block ERD is half a period of the frequency under investigation. (2) Due to the slow impulse response, amplitude effects in the ERD may in reality be caused by duration differences. (3) Although ERD based on the Hilbert transform does not yield any significant advantages over classical ERD in terms of time resolution, it has some important practical advantages.

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

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

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

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

In various applications of forensic phonetics the question arises as to how far aural-perceptual speaker recognition performance is reliable. Therefore, it is necessary to examine the relationship between speaker recognition results and human perception/production abilities like musicality or speech sensitivity. In this study, performance in a speaker recognition experiment and a speech sensitivity test are correlated. The results show a moderately significant positive correlation between the two tasks. Generally, performance in the speaker recognition task was better than in the speech sensitivity test. Professionals in speech and singing yielded a more homogeneous correlation than non-experts. Training in speech as well as choir-singing seems to have a positive effect on performance in speaker recognition. It may be concluded, firstly, that in cases where the reliability of voice line-up results or the credibility of a testimony have to be considered, the speech sensitivity test could be a useful indicator. Secondly, the speech sensitivity test might be integrated into the canon of possible procedures for the accreditation of forensic phoneticians. Both tests may also be used in combination.

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.