Publications

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

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

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

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

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

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

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

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

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

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

People learn language from their social environment. As individuals differ in their social networks, they might be exposed to input with different lexical distributions, and these might influence their linguistic representations and lexical choices. In this article we test the relation between linguistic performance and 3 social network properties that should influence input variability, namely, network size, network heterogeneity, and network density. In particular, we examine how these social network properties influence lexical prediction, lexical access, and lexical use. To do so, in Study 1, participants predicted how people of different ages would name pictures, and in Study 2 participants named the pictures themselves. In both studies, we examined how participants’ social network properties related to their performance. In Study 3, we ran simulations on norms we collected to see how age variability in one’s network influences the distribution of different names in the input. In all studies, network age heterogeneity influenced performance leading to better prediction, faster response times for difficult-to-name items, and less entropy in input distribution. These results suggest that individual differences in social network properties can influence linguistic behavior. Specifically, they show that having a more heterogeneous network is associated with better performance. These results also show that the same factors influence lexical prediction and lexical production, suggesting the two might be related.

Abstract

Speakers constantly learn language from the environment by sampling their linguistic input and adjusting their representations accordingly. Logically, people should attend more to the environment and adjust their behavior in accordance with it more the lower their success in the environment is. We test whether the learning of linguistic input follows this general principle in two studies: a corpus analysis of a TV game show, Jeopardy, and a laboratory task modeled after Go Fish. We show that lower (non-linguistic) success in the task modulates learning of and reliance on linguistic patterns in the environment. In Study 1, we find that poorer performance increases conformity with linguistic norms, as reflected by increased preference for frequent grammatical structures. In Study 2, which consists of a more interactive setting, poorer performance increases learning from the immediate social environment, as reflected by greater repetition of others’ grammatical structures. We propose that these results have implications for models of language production and language learning and for the propagation of language change. In particular, they suggest that linguistic changes might spread more quickly in times of crisis, or when the gap between more and less successful people is larger. The results might also suggest that innovations stem from successful individuals while their propagation would depend on relatively less successful individuals. We provide a few historical examples that are in line with the first suggested implication, namely, that the spread of linguistic changes is accelerated during difficult times, such as war time and an economic downturn

Abstract

Foreign-accented speech is generally harder to understand than native-accented speech. This difficulty is reduced for non-native listeners who share their first language with the non-native speaker. It is currently unclear, however, how non-native listeners deal with foreign-accented speech produced by speakers of a different language. We show that the process of (second) language acquisition is associated with an increase in the relative difficulty of processing foreign-accented speech. Therefore, experiencing greater relative difficulty with foreign-accented speech compared with native speech is a marker of language proficiency. These results contribute to our understanding of how phonological categories are acquired during second language learning.

Abstract

We learn language from our social environment. In general, the more sources we have, the less informative each of them is, and the less weight we should assign it. If this is the case, people who interact with fewer others should be more susceptible to the influence of each of their interlocutors. This paper tests whether indeed people who interact with fewer other people have more malleable phonological representations. Using a perceptual learning paradigm, this paper shows that individuals who regularly interact with fewer others are more likely to change their boundary between /d/ and /t/ following exposure to an atypical speaker. It further shows that the effect of number of interlocutors is not due to differences in ability to learn the speaker’s speech patterns, but specific to likelihood of generalizing the learned pattern. These results have implications for both language learning and language change, as they suggest that individuals with smaller social networks might play an important role in propagating linguistic changes.

Abstract

The order of acquisition of Dutch syllable types by first language learners is analyzed as following from an initial ranking and subsequent rerankings of constraints in an optimality theoretic grammar. Initially, structural constraints are all ranked above faithfulness constraints, leading to core syllable (CV) productions only. Subsequently, faithfulness gradually rises to the highest position in the ranking, allowing more and more marked syllable types to appear in production. Local conjunctions of Structural constraints allow for a more detailed analysis.

Abstract

Preparing words in speech production is normally a fast and accurate process. We generate them two or three per second in fluent conversation; and overtly naming a clear picture of an object can easily be initiated within 600 msec after picture onset. The underlying process, however, is exceedingly complex. The theory reviewed in this target article analyzes this process as staged and feedforward. After a first stage of conceptual preparation, word generation proceeds through lexical selection, morphological and phonological encoding, phonetic encoding, and articulation itself. In addition, the speaker exerts some degree of output control, by monitoring of self-produced internal and overt speech. The core of the theory, ranging from lexical selection to the initiation of phonetic encoding, is captured in a computational model, called WEAVER + +. Both the theory and the computational model have been developed in interaction with reaction time experiments, particularly in picture naming or related word production paradigms, with the aim of accounting. for the real-time processing in normal word production. A comprehensive review of theory, model, and experiments is presented. The model can handle some of the main observations in the domain of speech errors (the major empirical domain for most other theories of lexical access), and the theory opens new ways of approaching the cerebral organization of speech production by way of high-temporal-resolution imaging.

Abstract

The purpose of this study was to relate a psycholinguistic processing model of picture naming to the dynamics of cortical activation during picture naming. The activation was recorded from eight Dutch subjects with a whole-head neuromagnetometer. The processing model, based on extensive naming latency studies, is a stage model. In preparing a picture's name, the speaker performs a chain of specific operations. They are, in this order, computing the visual percept, activating an appropriate lexical concept, selecting the target word from the mental lexicon, phonological encoding, phonetic encoding, and initiation of articulation. The time windows for each of these operations are reasonably well known and could be related to the peak activity of dipole sources in the individual magnetic response patterns. The analyses showed a clear progression over these time windows from early occipital activation, via parietal and temporal to frontal activation. The major specific findings were that (1) a region in the left posterior temporal lobe, agreeing with the location of Wernicke's area, showed prominent activation starting about 200 msec after picture onset and peaking at about 350 msec, (i.e., within the stage of phonological encoding), and (2) a consistent activation was found in the right parietal cortex, peaking at about 230 msec after picture onset, thus preceding and partly overlapping with the left temporal response. An interpretation in terms of the management of visual attention is proposed.

Abstract

Research on spoken word production has been approached from two angles. In one research tradition, the analysis of spontaneous or induced speech errors led to models that can account for speech error distributions. In another tradition, the measurement of picture naming latencies led to chronometric models accounting for distributions of reaction times in word production. Both kinds of models are, however, dealing with the same underlying processes: (1) the speaker’s selection of a word that is semantically and syntactically appropriate; (2) the retrieval of the word’s phonological properties; (3) the rapid syllabification of the word in context; and (4) the preparation of the corresponding articulatory gestures. Models of both traditions explain these processes in terms of activation spreading through a localist, symbolic network. By and large, they share the main levels of representation: conceptual/semantic, syntactic, phonological and phonetic. They differ in various details, such as the amount of cascading and feedback in the network. These research traditions have begun to merge in recent years, leading to highly constructive experimentation. Currently, they are like two similar knives honing each other. A single pair of scissors is in the making.

Abstract

This commentary discusses whether abstract metrical frames are stored. For stress-assigning languages (e.g., Dutch and English), which have a dominant stress pattern, metrical frames are stored only for words that deviate from the default stress pattern. The majority of the words in these languages are produced without retrieving any independent syllabic or metrical frame.

Abstract

In their comment, Dell and O'Seaghdha (1991) adduced any effect on phonological probes for semantic alternatives to the activation of these probes in the lexical network. We argue that that interpretation is false and, in addition, that the model still cannot account for our data. Furthermore, and different from Dell and O'seaghda, we adduce semantic rebound to the lemma level, where it is so substantial that it should have shown up in our data. Finally, we question the function of feedback in a lexical network (other than eliciting speech errors) and discuss Dell's (1988) notion of a unified production-comprehension system.

Abstract

In recent decades, psychologists have become increasingly interested in our ability to speak. This paper sketches the present theoretical perspective on this most complex skill of homo sapiens. The generation of fluent speech is based on the interaction of various processing components. These mechanisms are highly specialized, dedicated to performing specific subroutines, such as retrieving appropriate words, generating morpho-syntactic structure, computing the phonological target shape of syllables, words, phrases and whole utterances, and creating and executing articulatory programmes. As in any complex skill, there is a self-monitoring mechanism that checks the output. These component processes are targets of increasingly sophisticated experimental research, of which this paper presents a few salient examples.

Abstract

The core issue in the 19-century sources of psycholinguistics was the question, "Where does language come from?'' This genetic perspective unified the study of the ontogenesis, the phylogenesis, the microgenesis, and to some extent the neurogenesis of language. This paper makes the point that this original perspective is still a valid and attractive one. It is exemplified by a discussion of the genesis of spoken words.

Abstract

It has been hypothesised that human reasoning has a non-linguistic foundation, but is nevertheless influenced by the formal means available in a language. For example, Western logic is transparently related to European sentential connectives (e.g., and, if … then, or, not), some of which cannot be unambiguously expressed in other languages. The questionnaire explores reasoning tools and practices through investigating translation equivalents of English sentential connectives and collecting examples of “reasoned arguments”.

Abstract

Demonstratives are key items in understanding how a language constructs and interprets spatial relationships. They are also multi-functional, with applications to non-spatial deictic fields such as time, perception, person and discourse, and uses in anaphora and affect marking. This item consists of an overview of theoretical distinctions in demonstrative systems, followed by a set of practical queries and elicitation suggestions for demonstratives in “table top” space, wider spatial fields, and naturalistic data.

Abstract

The world’s languages encode a diverse range of topological relations. However, cross-linguistic investigation suggests that the relations IN, AT and ON are especially fundamental to the grammaticised expression of space. The purpose of this questionnaire is to collect information about adpositions, case markers, and spatial nominals that are involved in the expression of core IN/AT/ON meanings. The task explores the more general parts of a language’s topological system, with a view to testing certain hypotheses about the packaging of spatial concepts. The questionnaire consists of target translation sentences that focus on a number of dimensions including animacy, caused location and motion.

Abstract

Languages differ widely in terms of how they encode the fundamental concepts of location and position. For some languages, verbs have an important role to play in describing situations (e.g., whether a bottle is standing or lying on the table); for others, verbs are not used in describing location at all. This item outlines certain hypotheses concerning four “types” of languages: those that have verbless basic locatives; those that use a single verb; those that have several verbs available to express location; and those that use positional verbs. The document was originally published as an appendix to the 'Picture series for positional verbs' (https://doi.org/10.17617/2.2573831).

Abstract

This review describes some recent, unexpected findings concerning variation in spatial language across cultures, and places them in the context of the general anthropology of space on the one hand, and theories of spatial cognition in the cognitive sciences on the other. There has been much concern with the symbolism of space in anthropological writings, but little on concepts of space in practical activities. This neglect of everyday spatial notions may be due to unwitting ethnocentrism, the assumption in Western thinking generally that notions of space are universally of a single kind. Recent work shows that systems of spatial reckoning and description can in fact be quite divergent across cultures, linguistic differences correlating with distinct cognitive tendencies. This unexpected cultural variation raises interesting questions concerning the relation between cultural and linguistic concepts and the biological foundations of cognition. It argues for more sophisticated models relating culture and cognition than we currently have available.