Publications

Abstract

Using eye-tracking as a window on cognitive processing, this study investigates language effects on attention to motion events in a non-verbal task. We compare gaze allocation patterns by native speakers of German and Modern Standard Arabic (MSA), two languages that differ with regard to the grammaticalization of temporal concepts. Findings of the non-verbal task, in which speakers watch dynamic event scenes while performing an auditory distracter task, are compared to gaze allocation patterns which were obtained in an event description task, using the same stimuli. We investigate whether differences in the grammatical aspectual systems of German and MSA affect the extent to which endpoints of motion events are linguistically encoded and visually processed in the two tasks. In the linguistic task, we find clear language differences in endpoint encoding and in the eye-tracking data (attention to event endpoints) as well: German speakers attend to and linguistically encode endpoints more frequently than speakers of MSA. The fixation data in the non-verbal task show similar language effects, providing relevant insights with regard to the language-and-thought debate. The present study is one of the few studies that focus explicitly on language effects related to grammatical concepts, as opposed to lexical concepts.

Abstract

In well-known demonstrations of lexical prediction during language comprehension, pre-nominal articles that mismatch a likely upcoming noun's gender elicit different neural activity than matching articles. However, theories differ on what this pre-nominal prediction effect means and on what is being predicted. Does it reflect mismatch with a predicted article, or ‘merely’ revision of the noun prediction? We contrasted the ‘article prediction mismatch’ hypothesis and the ‘noun prediction revision’ hypothesis in two ERP experiments on Dutch mini-story comprehension, with pre-registered data collection and analyses. We capitalized on the Dutch gender system, which marks gender on definite articles (‘de/het’) but not on indefinite articles (‘een’). If articles themselves are predicted, mismatching gender should have little effect when readers expected an indefinite article without gender marking. Participants read contexts that strongly suggested either a definite or indefinite noun phrase as its best continuation, followed by a definite noun phrase with the expected noun or an unexpected, different gender noun phrase (‘het boek/de roman’, the book/the novel). Experiment 1 (N = 48) showed a pre-nominal prediction effect, but evidence for the article prediction mismatch hypothesis was inconclusive. Informed by exploratory analyses and power analyses, direct replication Experiment 2 (N = 80) yielded evidence for article prediction mismatch at a newly pre-registered occipital region-of-interest. However, at frontal and posterior channels, unexpectedly definite articles also elicited a gender-mismatch effect, and this support for the noun prediction revision hypothesis was further strengthened by exploratory analyses: ERPs elicited by gender-mismatching articles correlated with incurred constraint towards a new noun (next-word entropy), and N400s for initially unpredictable nouns decreased when articles made them more predictable. By demonstrating its dual nature, our results reconcile two prevalent explanations of the pre-nominal prediction effect.

Abstract

In this event-related FMRI study we investigated the effect of five days of implicit acquisition on preference classification by means of an artificial grammar learning (AGL) paradigm based on the structural mere-exposure effect and preference classification using a simple right-linear unification grammar. This allowed us to investigate implicit AGL in a proper learning design by including baseline measurements prior to grammar exposure. After 5 days of implicit acquisition, the FMRI results showed activations in a network of brain regions including the inferior frontal (centered on BA 44/45) and the medial prefrontal regions (centered on BA 8/32). Importantly, and central to this study, the inclusion of a naive preference FMRI baseline measurement allowed us to conclude that these FMRI findings were the intrinsic outcomes of the learning process itself and not a reflection of a preexisting functionality recruited during classification, independent of acquisition. Support for the implicit nature of the knowledge utilized during preference classification on day 5 come from the fact that the basal ganglia, associated with implicit procedural learning, were activated during classification, while the medial temporal lobe system, associated with explicit declarative memory, was consistently deactivated. Thus, preference classification in combination with structural mere-exposure can be used to investigate structural sequence processing (syntax) in unsupervised AGL paradigms with proper learning designs.

Abstract

Objective: To combine MRI-based cortical morphometry and diffusion white matter tractography to describe the anatomical correlates of repetition deficits in patients with primary progressive aphasia (PPA).

Methods: The traditional anatomical model of language identifies a network for word repetition that includes Wernicke and Broca regions directly connected via the arcuate fasciculus. Recent tractography findings of an indirect pathway between Wernicke and Broca regions suggest a critical role of the inferior parietal lobe for repetition. To test whether repetition deficits are associated with damage to the direct or indirect pathway between both regions, tractography analysis was performed in 30 patients with PPA (64.27 ± 8.51 years) and 22 healthy controls. Cortical volume measurements were also extracted from 8 perisylvian language areas connected by the direct and indirect pathways.

Results: Compared to healthy controls, patients with PPA presented with reduced performance in repetition tasks and increased damage to most of the perisylvian cortical regions and their connections through the indirect pathway. Repetition deficits were prominent in patients with cortical atrophy of the temporo-parietal region with volumetric reductions of the indirect pathway.

Conclusions: The results suggest that in PPA, deficits in repetition are due to damage to the temporo-parietal cortex and its connections to Wernicke and Broca regions. We therefore propose a revised language model that also includes an indirect pathway for repetition, which has important clinical implications for the functional mapping and treatment of neurologic patients.

Abstract

Stroke-induced aphasia is associated with adverse effects on quality of life and the ability to return to work. For patients and clinicians the possibility of relying on valid predictors of recovery is an important asset in the clinical management of stroke-related impairment. Age, level of education, type and severity of initial symptoms are established predictors of recovery. However, anatomical predictors are still poorly understood. In this prospective longitudinal study, we intended to assess anatomical predictors of recovery derived from diffusion tractography of the perisylvian language networks. Our study focused on the arcuate fasciculus, a language pathway composed of three segments connecting Wernicke’s to Broca’s region (i.e. long segment), Wernicke’s to Geschwind’s region (i.e. posterior segment) and Broca’s to Geschwind’s region (i.e. anterior segment). In our study we were particularly interested in understanding how lateralization of the arcuate fasciculus impacts on severity of symptoms and their recovery. Sixteen patients (10 males; mean age 60 ± 17 years, range 28–87 years) underwent post stroke language assessment with the Revised Western Aphasia Battery and neuroimaging scanning within a fortnight from symptoms onset. Language assessment was repeated at 6 months. Backward elimination analysis identified a subset of predictor variables (age, sex, lesion size) to be introduced to further regression analyses. A hierarchical regression was conducted with the longitudinal aphasia severity as the dependent variable. The first model included the subset of variables as previously defined. The second model additionally introduced the left and right arcuate fasciculus (separate analysis for each segment). Lesion size was identified as the only independent predictor of longitudinal aphasia severity in the left hemisphere [beta = −0.630, t(−3.129), P = 0.011]. For the right hemisphere, age [beta = −0.678, t(–3.087), P = 0.010] and volume of the long segment of the arcuate fasciculus [beta = 0.730, t(2.732), P = 0.020] were predictors of longitudinal aphasia severity. Adding the volume of the right long segment to the first-level model increased the overall predictive power of the model from 28% to 57% [F(1,11) = 7.46, P = 0.02]. These findings suggest that different predictors of recovery are at play in the left and right hemisphere. The right hemisphere language network seems to be important in aphasia recovery after left hemispheric stroke.

Abstract

The occipital and frontal lobes are anatomically distant yet functionally highly integrated to generate some of the most complex behaviour. A series of long associative fibres, such as the fronto-occipital networks, mediate this integration via rapid feed-forward propagation of visual input to anterior frontal regions and direct top–down modulation of early visual processing.

Despite the vast number of anatomical investigations a general consensus on the anatomy of fronto-occipital connections is not forthcoming. For example, in the monkey the existence of a human equivalent of the ‘inferior fronto-occipital fasciculus’ (iFOF) has not been demonstrated. Conversely, a ‘superior fronto-occipital fasciculus’ (sFOF), also referred to as ‘subcallosal bundle’ by some authors, is reported in monkey axonal tracing studies but not in human dissections.

In this study our aim is twofold. First, we use diffusion tractography to delineate the in vivo anatomy of the sFOF and the iFOF in 30 healthy subjects and three acallosal brains. Second, we provide a comprehensive review of the post-mortem and neuroimaging studies of the fronto-occipital connections published over the last two centuries, together with the first integral translation of Onufrowicz's original description of a human fronto-occipital fasciculus (1887) and Muratoff's report of the ‘subcallosal bundle’ in animals (1893).

Our tractography dissections suggest that in the human brain (i) the iFOF is a bilateral association pathway connecting ventro-medial occipital cortex to orbital and polar frontal cortex, (ii) the sFOF overlaps with branches of the superior longitudinal fasciculus (SLF) and probably represents an ‘occipital extension’ of the SLF, (iii) the subcallosal bundle of Muratoff is probably a complex tract encompassing ascending thalamo-frontal and descending fronto-caudate connections and is therefore a projection rather than an associative tract.

In conclusion, our experimental findings and review of the literature suggest that a ventral pathway in humans, namely the iFOF, mediates a direct communication between occipital and frontal lobes. Whether the iFOF represents a unique human pathway awaits further ad hoc investigations in animals.

Abstract

This scientific commentary refers to ‘Metabolic lesion-deficit mapping of human cognition’ by Jha etal.

Abstract

In speech, listeners extract continuously-varying spectrotemporal cues from the acoustic signal to perceive discrete phonetic categories. Spectral cues are spatially encoded in the amplitude of responses in phonetically-tuned neural populations in auditory cortex. It remains unknown whether similar neurophysiological mechanisms encode temporal cues like voice-onset time (VOT), which distinguishes sounds like /b/ and/p/. We used direct brain recordings in humans to investigate the neural encoding of temporal speech cues with a VOT continuum from /ba/ to /pa/. We found that distinct neural populations respond preferentially to VOTs from one phonetic category, and are also sensitive to sub-phonetic VOT differences within a population’s preferred category. In a simple neural network model, simulated populations tuned to detect either temporal gaps or coincidences between spectral cues captured encoding patterns observed in real neural data. These results demonstrate that a spatial/amplitude neural code underlies the cortical representation of both spectral and temporal speech cues.

Abstract

Vocabulary learning occurs throughout the lifespan, often implicitly. For foreign language learners,
this is particularly challenging as they must acquire a large number of new words with little exposure.
In the present study, we explore the effects of contextual diversity—namely, the number of texts a
word appears in—on native and foreign language word learning. Participants read several texts that
had novel pseudowords replacing high-frequency words. The total number of encounters with the
novel words was held constant, but they appeared in 1, 2, 4, or 8 texts. In addition, some participants
read the texts in Spanish (their native language) and others in English (their foreign language). We
found that increasing contextual diversity improved recall and recognition of the word, as well as the
ability to match the word with its meaning while keeping comprehension unimpaired. Using a foreign
language only affected performance in the matching task, where participants had to quickly identify
the meaning of the word. Results are discussed in the greater context of the word learning and foreign
language literature as well as their importance as a teaching tool.

Abstract

The current study focuses on how different scales with varying demands can
affect our subjective assessments. We carried out 2 experiments in which we
asked participants to rate how happy or sad morphed images of faces looked.
The two extremes were the original happy and original sad faces with 4
morphs in between. We manipulated language of the task—namely, half of
the participants carried it out in their native language, Spanish, and the other
half in their foreign language, English—and type of scale. Within type of
scale, we compared verbal and brightness scales. We found that, while
language did not have an effect on the assessment, type of scale did. The
brightness scale led to overall higher ratings, i.e., assessing all faces as
somewhat happier. This provides a limitation on the foreign language effect,
as well as evidence for the influence of the cognitive demands of a scale on
emotionality assessments.

Abstract

Learning new content and vocabulary in a foreign language can be particularly difficult. Yet,
there are educational programs that require people to study in a language they are not
native speakers of. For this reason, it is important to understand how these learning processes work and possibly differ from native language learning, as well as to develop strategies to ease this process. The current study takes advantage of emotionality—operationally
defined as positive valence and high arousal—to improve memory. In two experiments, the
present paper addresses whether participants have more difficulty learning the names of
objects they have never seen before in their foreign language and whether embedding them
in a positive semantic context can help make learning easier. With this in mind, we had participants (with a minimum of a B2 level of English) in two experiments (43 participants in
Experiment 1 and 54 in Experiment 2) read descriptions of made-up objects—either positive
or neutral and either in their native or a foreign language. The effects of language varied
with the difficulty of the task and measure used. In both cases, learning the words in a positive context improved learning. Importantly, the effect of emotionality was not modulated by
language, suggesting that the effects of emotionality are independent of language and could
potentially be a useful tool for improving foreign language vocabulary learning.

Abstract

Previous research on the effect of perturbed auditory feedback in speech production has focused on two types of responses. In the short term, speakers generate compensatory motor commands in response to unexpected perturbations. In the longer term, speakers adapt feedforward motor programmes in response to feedback perturbations, to avoid future errors. The current study investigated the relation between these two types of responses to altered auditory feedback. Specifically, it was hypothesised that consistency in previous feedback perturbations would influence whether speakers adapt their feedforward motor programmes. In an altered auditory feedback paradigm, formant perturbations were applied either across all trials (the consistent condition) or only to some trials, whereas the others remained unperturbed (the inconsistent condition). The results showed that speakers’ responses were affected by feedback consistency, with stronger speech changes in the consistent condition compared with the inconsistent condition. Current models of speech-motor control can explain this consistency effect. However, the data also suggest that compensation and adaptation are distinct processes, which are not in line with all current models.

Abstract

Kleefstra syndrome (KS) is a neurodevelopmental disorder caused by mutations in the histone methyltransferase EHMT1. To study the impact of decreased EHMT1 function in human cells, we generated excitatory cortical neurons from induced pluripotent stem (iPS) cells derived from KS patients. Neuronal networks of patient-derived cells exhibit network bursting with a reduced rate, longer duration, and increased temporal irregularity compared to control networks. We show that these changes are mediated by upregulation of NMDA receptor (NMDAR) subunit 1 correlating with reduced deposition of the repressive H3K9me2 mark, the catalytic product of EHMT1, at the GRIN1 promoter. In mice EHMT1 deficiency leads to similar neuronal network impairments with increased NMDAR function. Finally, we rescue the KS patient-derived neuronal network phenotypes by pharmacological inhibition of NMDARs. Summarized, we demonstrate a direct link between EHMT1 deficiency and NMDAR hyperfunction in human neurons, providing a potential basis for more targeted therapeutic approaches for KS.

Abstract

Disruptions of the FOXP2 gene cause a speech and language disorder involving difficulties in sequencing orofacial movements. FOXP2 is expressed in cortico-striatal and cortico-cerebellar circuits important for fine motor skills, and affected individuals show abnormalities in these brain regions. We selectively disrupted Foxp2 in the cerebellar Purkinje cells, striatum or cortex of mice and assessed the effects on skilled motor behaviour using an operant lever-pressing task. Foxp2 loss in each region impacted behaviour differently, with striatal and Purkinje cell disruptions affecting the variability and the speed of lever-press sequences, respectively. Mice lacking Foxp2 in Purkinje cells showed a prominent phenotype involving slowed lever pressing as well as deficits in skilled locomotion. In vivo recordings from Purkinje cells uncovered an increased simple spike firing rate and decreased modulation of firing during limb movements. This was caused by increased intrinsic excitability rather than changes in excitatory or inhibitory inputs. Our findings show that Foxp2 can modulate different aspects of motor behaviour in distinct brain regions, and uncover an unknown role for Foxp2 in the modulation of Purkinje cell activity that severely impacts skilled movements.

Abstract

Disruptions of the FOXP2 gene cause a rare speech and language disorder, a discovery that has opened up novel avenues for investigating the relevant neural pathways. FOXP2 shows remarkably high conservation of sequence and neural expression in diverse vertebrates, suggesting that studies in other species are useful in elucidating its functions. Here we describe how investigations of mice that carry disruptions of Foxp2 provide insights at multiple levels: molecules, cells, circuits and behaviour. Work thus far has implicated the gene in key processes including neurite outgrowth, synaptic plasticity, sensorimotor integration and motor-skill learning.

Abstract

Human behavioral asymmetries are commonly studied in the context of structural cortical and connectional asymmetries. Within this framework, Sreenivasan and Sridharan (1) provide intriguing evidence of a relationship between visual asymmetries and the lateralization of superior colliculi connections—a phylogenetically older mesencephalic structure. Specifically, response facilitation for cued locations (i.e., choice bias) in the contralateral hemifield was associated with differences in the connectivity of the superior colliculus. Given that the superior colliculus has a structural homolog—the optic tectum—which can be traced across all Vertebrata, these results may have meaningful evolutionary ramifications.

Abstract

Gradients capture some of the variance of the resting-state functional magnetic resonance imaging (rsfMRI) signal. Amongst these, the principal gradient depicts a functional processing hierarchy that spans from sensory-motor cortices to regions of the default-mode network. While the cortex has been well characterised in terms of gradients little is known about its underlying white matter. For instance, comprehensive mapping of the principal gradient on the largest white matter tract, the corpus callosum, is still missing. Here, we mapped the principal gradient onto the midsection of the corpus callosum using the 7T human connectome project dataset. We further explored how quantitative measures and variability in callosal midsection connectivity relate to the principal gradient values. In so doing, we demonstrated that the extreme values of the principal gradient are located within the callosal genu and the posterior body, have lower connectivity variability but a larger spatial extent along the midsection of the corpus callosum than mid-range values. Our results shed light on the relationship between the brain's functional hierarchy and the corpus callosum. We further speculate about how these results may bridge the gap between functional hierarchy, brain asymmetries, and evolution.

Abstract

High frequency words have been suggested to benefit both speech segmentation and grammatical categorization of the words around them. Despite utilizing similar information, these tasks are usually investigated separately in studies examining learning. We determined whether including high frequency words in continuous speech could support categorization when words are being segmented for the first time. We familiarized learners with continuous artificial speech comprising repetitions of target words, which were preceded by high-frequency marker words. Crucially, marker words distinguished targets into 2 distributionally defined categories. We measured learning with segmentation and categorization tests and compared performance against a control group that heard the artificial speech without these marker words (i.e., just the targets, with no cues for categorization). Participants segmented the target words from speech in both conditions, but critically when the marker words were present, they influenced acquisition of word-referent mappings in a subsequent transfer task, with participants demonstrating better early learning for mappings that were consistent (rather than inconsistent) with the distributional categories. We propose that high-frequency words may assist early grammatical categorization, while speech segmentation is still being learned.

Abstract

High frequency words play a key role in language acquisition, with recent work suggesting they may serve both speech segmentation and lexical categorisation. However, it is not yet known whether infants can detect novel high frequency words in continuous speech, nor whether they can use them to help learning for segmentation and categorisation at the same time. For instance, when hearing “you eat the biscuit”, can children use the high-frequency words “you” and “the” to segment out “eat” and “biscuit”, and determine their respective lexical categories? We tested this in two experiments. In Experiment 1, we familiarised 12-month-old infants with continuous artificial speech comprising repetitions of target words, which were preceded by high-frequency marker words that distinguished the targets into two distributional categories. In Experiment 2, we repeated the task using the same language but with additional phonological cues to word and category structure. In both studies, we measured learning with head-turn preference tests of segmentation and categorisation, and compared performance against a control group that heard the artificial speech without the marker words (i.e., just the targets). There was no evidence that high frequency words helped either speech segmentation or grammatical categorisation. However, segmentation was seen to improve when the distributional information was supplemented with phonological cues (Experiment 2). In both experiments, exploratory analysis indicated that infants’ looking behaviour was related to their linguistic maturity (indexed by infants’ vocabulary scores) with infants with high versus low vocabulary scores displaying novelty and familiarity preferences, respectively. We propose that high-frequency words must reach a critical threshold of familiarity before they can be of significant benefit to learning.

Abstract

To acquire language, infants must learn how to identify words and linguistic structure in speech. Statistical learning has been suggested to assist both of these tasks. However, infants’ capacity to use statistics to discover words and structure together remains unclear. Further, it is not yet known how infants’ statistical learning ability relates to their language development. We trained 17-month-old infants on an artificial language comprising non-adjacent dependencies, and examined their looking times on tasks assessing sensitivity to words and structure using an eye-tracked head-turn-preference paradigm. We measured infants’ vocabulary size using a Communicative Development Inventory (CDI) concurrently and at 19, 21, 24, 25, 27, and 30 months to relate performance to language development. Infants could segment the words from speech, demonstrated by a significant difference in looking times to words versus part-words. Infants’ segmentation performance was significantly related to their vocabulary size (receptive and expressive) both currently, and over time (receptive until 24 months, expressive until 30 months), but was not related to the rate of vocabulary growth. The data also suggest infants may have developed sensitivity to generalised structure, indicating similar statistical learning mechanisms may contribute to the discovery of words and structure in speech, but this was not related to vocabulary size.

Abstract

Cumulative tool-based culture underwrote our species' evolutionary success and tool-based nut-cracking is one of the strongest candidates for cultural transmission in our closest relatives, chimpanzees. However the social learning processes that may explain both the similarities and differences between the species remain unclear. A previous study of nut-cracking by initially naïve chimpanzees suggested that a learning chimpanzee holding no hammer nevertheless replicated hammering actions it witnessed. This observation has potentially important implications for the nature of the social learning processes and underlying motor coding involved. In the present study, model and observer actions were quantified frame-by-frame and analysed with stringent statistical methods, demonstrating synchrony between the observer's and model's movements, cross-correlation of these movements above chance level and a unidirectional transmission process from model to observer. These results provide the first quantitative evidence for motor mimicking underlain by motor coding in apes, with implications for mirror neuron function.

Abstract

Previous research on language development shows that children are tuned early on to the language-specific semantic and syntactic encoding of events in their native language. Here we ask whether language-specificity is also evident in children's early representations in gesture accompanying speech. In a longitudinal study, we examined the spontaneous speech and cospeech gestures of eight Turkish-speaking children aged one to three and focused on their caused motion event expressions. In Turkish, unlike in English, the main semantic elements of caused motion such as Action and Path can be encoded in the verb (e.g. sok- ‘put in’) and the arguments of a verb can be easily omitted. We found that Turkish-speaking children's speech indeed displayed these language-specific features and focused on verbs to encode caused motion. More interestingly, we found that their early gestures also manifested specificity. Children used iconic cospeech gestures (from 19 months onwards) as often as pointing gestures and represented semantic elements such as Action with Figure and/or Path that reinforced or supplemented speech in language-specific ways until the age of three. In the light of previous reports on the scarcity of iconic gestures in English-speaking children's early productions, we argue that the language children learn shapes gestures and how they get integrated with speech in the first three years of life.

Abstract

Background: Two distinct insomnia disorder (ID) phenotypes have been proposed, distinguished on the basis of an objective total sleep time less or more than 6 hr. In particular, it has been recently reported that patients with objective short sleep duration have a blunted response to cognitive behavioral therapy for insomnia (CBT-I). The aim of this study was to investigate the differences of CBT-I response in two groups of ID patients subdivided according to total sleep time. Methods: Two hundred forty-six ID patients were subdivided into two groups, depending on their reported total sleep time (TST) assessed by sleep diaries. Patients with a TST greater than 6 hr were classified as “normal sleepers” (NS), while those with a total sleep time less than 6 hr were classified as “short sleepers” (SS). Results: The delta between Insomnia Severity Index scores and sleep efficiency at the beginning as compared to the end of the treatment was significantly higher for SS in comparison to NS, even if they still exhibit more insomnia symptoms. No difference was found between groups in terms of remitters; however, more responders were observed in the SS group in comparison to the NS group. Conclusions: Our results demonstrate that ID patients with reported short total sleep time had a beneficial response to CBT-I of greater magnitude in comparison to NS. However, these patients may still experience the presence of residual insomnia symptoms after treatment.

Abstract

Several studies report an association between REM Sleep Behavior Disorder (RBD) and neurodegenerative diseases, in particular synucleinopathies. Interestingly, the onset of RBD precedes the development of neurodegeneration by several years. This review and meta-analysis aims to establish the rate of conversion of RBD into neurodegenerative diseases. Longitudinal studies were searched from the PubMed, Web of Science, and SCOPUS databases. Using random-effect modeling, we performed a meta-analysis on the rate of RBD conversions into neurodegeneration. Furthermore, we fitted a Kaplan-Meier analysis and compared the differences between survival curves of different diseases with log-rank tests. The risk for developing neurodegenerative diseases was 33.5% at five years follow-up, 82.4% at 10.5 years and 96.6% at 14 years. The average conversion rate was 31.95% after a mean duration of follow-up of 4.75 ± 2.43 years. The majority of RBD patients converted to Parkinson's Disease (43%), followed by Dementia with Lewy Bodies (25%). The estimated risk for RBD patients to develop a neurodegenerative disease over a long-term follow-up is more than 90%. Future studies should include control group for the evaluation of REM sleep without atonia as marker for neurodegeneration also in non-clinical population and target RBD as precursor of neurodegeneration to develop protective trials.

Abstract

Visuospatial attention theories often propose hemispheric asymmetries underlying the control of attention. In general support of these theories, previous EEG/MEG studies have shown that spatial attention is associated with hemispheric modulation of posterior alpha power (gating by inhibition). However, since measures of alpha power are typically expressed as lateralization scores, or collapsed across left and right attention shifts, the individual hemispheric contribution to the attentional control mechanism remains unclear. This is, however, the most crucial and decisive aspect in which the currently competing attention theories continue to disagree. To resolve this long-standing conflict, we derived predictions regarding alpha power modulations from Heilman's hemispatial theory and Kinsbourne's interhemispheric competition theory and tested them empirically in an EEG experiment. We used an attention paradigm capable of isolating alpha power modulation in two attentional states, namely attentional bias in a neutral cue condition and spatial orienting following directional cues. Differential alpha modulations were found for both hemispheres across conditions. When anticipating peripheral visual targets without preceding directional cues (neutral condition), posterior alpha power in the left hemisphere was generally lower and more strongly modulated than in the right hemisphere, in line with the interhemispheric competition theory. Intriguingly, however, while alpha power in the right hemisphere was modulated by both, cue-directed leftward and rightward attention shifts, the left hemisphere only showed modulations by rightward shifts of spatial attention, in line with the hemispatial theory. This suggests that the two theories may not be mutually exclusive, but rather apply to different attentional states.

Abstract

This study investigated how sentence formulation is influenced by a preceding discourse context. In two eye-tracking experiments, participants described pictures of two-character transitive events in Dutch (Experiment 1) and Chinese (Experiment 2). Focus was manipulated by presenting questions before each picture. In the Neutral condition, participants first heard ‘What is happening here?’ In the Object or Subject Focus conditions, the questions asked about the Object or Subject character (What is the policeman stopping? Who is stopping the truck?). The target response was the same in all conditions (The policeman is stopping the truck). In both experiments, sentence formulation in the Neutral condition showed the expected pattern of speakers fixating the subject character (policeman) before the object character (truck). In contrast, in the focus conditions speakers rapidly directed their gaze preferentially only to the character they needed to encode to answer the question (the new, or focused, character). The timing of gaze shifts to the new character varied by language group (Dutch vs. Chinese): shifts to the new character occurred earlier when information in the question can be repeated in the response with the same syntactic structure (in Chinese but not in Dutch). The results show that discourse affects the timecourse of linguistic formulation in simple sentences and that these effects can be modulated by language-specific linguistic structures such as parallels in the syntax of questions and declarative sentences.

Abstract

Researchers have long avoided neurophysiological experiments of overt speech production due to the suspicion that artifacts caused by muscle activity may lead to a bad signal-to-noise ratio in the measurements. However, the need to actually produce speech may influence earlier processing and qualitatively change speech production processes and what we can infer from neurophysiological measures thereof. Recently, however, overt speech has been successfully investigated using EEG, MEG, and fMRI. The aim of this Research Topic is to draw together recent research on the neurophysiological basis of language production, with the aim of developing and extending theoretical accounts of the language production process. In this Research Topic of Frontiers in Language Sciences, we invite both experimental and review papers, as well as those about the latest methods in acquisition and analysis of overt language production data. All aspects of language production are welcome: i.e., from conceptualization to articulation during native as well as multilingual language production. Focus should be placed on using the neurophysiological data to inform questions about the processing stages of language production. In addition, emphasis should be placed on the extent to which the identified components of the electrophysiological signal (e.g., ERP/ERF, neuronal oscillations, etc.), brain areas or networks are related to language comprehension and other cognitive domains. By bringing together electrophysiological and neuroimaging evidence on language production mechanisms, a more complete picture of the locus of language production processes and their temporal and neurophysiological signatures will emerge.

Abstract

Learning to read a second language (L2) can pose a great challenge for children who have already been struggling to read in their first language (L1). Moreover, it is not clear whether, to what extent, and under what circumstances L1 reading difficulty increases the risk of L2 reading difficulty. This study investigated Chinese (L1) and English (L2) reading skills in a large representative sample of 1,824 Chinese–English bilingual children in Grades 4 and 5 from both urban and rural schools in Beijing. We examined the prevalence of reading difficulty in Chinese only (poor Chinese readers, PC), English only (poor English readers, PE), and both Chinese and English (poor bilingual readers, PB) and calculated the co-occurrence, that is, the chances of becoming a poor reader in English given that the child was already a poor reader in Chinese. We then conducted a multinomial logistic regression analysis and compared the prevalence of PC, PE, and PB between children in Grade 4 versus Grade 5, in urban versus rural areas, and in boys versus girls. Results showed that compared to girls, boys demonstrated significantly higher risk of PC, PE, and PB. Meanwhile, compared to the 5th graders, the 4th graders demonstrated significantly higher risk of PC and PB. In addition, children enrolled in the urban schools were more likely to become better second language readers, thus leading to a concerning rural–urban gap in the prevalence of L2 reading difficulty. Finally, among these Chinese–English bilingual children, regardless of sex and school location, poor reading skill in Chinese significantly increased the risk of also being a poor English reader, with a considerable and stable co-occurrence of approximately 36%. In sum, this study suggests that despite striking differences between alphabetic and logographic writing systems, L1 reading difficulty still significantly increases the risk of L2 reading difficulty. This indicates the shared meta-linguistic skills in reading different writing systems and the importance of understanding the universality and the interdependent relationship of reading between different writing systems. Furthermore, the male disadvantage (in both L1 and L2) and the urban–rural gap (in L2) found in the prevalence of reading difficulty calls for special attention to disadvantaged populations in educational practice.

Abstract

Previous research shows conflicting findings for the effect of font readability on comprehension and memory for language. It has been found that—perhaps counterintuitively–a hard to read font can be beneficial for language comprehension, especially for difficult language. Here we test how font readability influences the subjective experience of poetry reading. In three experiments we tested the influence of poem difficulty and font readability on the subjective experience of poems. We specifically predicted that font readability would have opposite effects on the subjective experience of easy versus difficult poems. Participants read poems which could be more or less difficult in terms of conceptual or structural aspects, and which were presented in a font that was either easy or more difficult to read. Participants read existing poems and subsequently rated their subjective experience (measured through four dependent variables: overall liking, perceived flow of the poem, perceived topic clarity, and perceived structure). In line with previous literature we observed a Poem Difficulty x Font Readability interaction effect for subjective measures of poetry reading. We found that participants rated easy poems as nicer when presented in an easy to read font, as compared to when presented in a hard to read font. Despite the presence of the interaction effect, we did not observe the predicted opposite effect for more difficult poems. We conclude that font readability can influence reading of easy and more difficult poems differentially, with strongest effects for easy poems.

Abstract

We investigated whether Tagalog-speaking children incrementally interpret the first noun
as the agent, even if verbal and nominal markers for assigning thematic roles are given
early in Tagalog sentences. We asked five- and seven-year-old children and adult
controls to select which of two pictures of reversible actions matched the sentence they
heard, while their looks to the pictures were tracked. Accuracy and eye-tracking data
showed that agent-initial sentences were easier to comprehend than patient-initial
sentences, but the effect of word order was modulated by voice. Moreover, our eyetracking
data provided evidence that, by the first noun phrase, seven-year-old children
looked more to the target in the agent-initial compared to the patient-initial conditions,
but this word order advantage was no longer observed by the second noun phrase. The
findings support language processing and acquisition models which emphasize the role
of frequency in developing heuristic strategies (e.g., Chang, Dell, & Bock, 2006).

Abstract

We report on two experiments that investigated the acquisition of the Tagalog symmetrical voice system, a typologically rare feature of Western Austronesian languages in which there are more than one basic transitive construction and no preference for agents to be syntactic subjects. In the experiments, 3-, 5-, and 7-year-old Tagalog-speaking children and adults completed a structural priming task that manipulated voice and word order, with the uniqueness of Tagalog allowing us to tease apart priming of thematic role order from that of syntactic roles. Participants heard a description of a picture showing a transitive action, and were then asked to complete a sentence of an unrelated picture using a voice-marked verb provided by the experimenter. Our results show that children gradually acquire an agent-before-patient preference, instead of having a default mapping of the agent to the first noun position. We also found an earlier mastery of the patient voice verbal and nominal marker configuration (patient is the subject), suggesting that children do not initially map the agent to the subject. Children were primed by thematic role but not syntactic role order, suggesting that they prioritize mapping of the thematic roles to sentence positions.

Abstract

Acoustic allometry is the study of how animal vocalisations reflect their body size. A key aim of this research is to identify outliers to acoustic allometry principles and pinpoint the evolutionary origins of such outliers. A parallel strand of research investigates species capable of vocal learning, the experience-driven ability to produce novel vocal signals through imitation or modification of existing vocalisations. Modification of vocalizations is a common feature found when studying both acoustic allometry and vocal learning. Yet, these two fields have only been investigated separately to date. Here, we review and connect acoustic allometry and vocal learning across mammalian clades, combining perspectives from bioacoustics, anatomy and evolutionary biology. Based on this, we hypothesize that, as a precursor to vocal learning, some species might have evolved the capacity for volitional vocal modulation via sexual selection for ‘dishonest’ signalling. We provide preliminary support for our hypothesis by showing significant associations between allometric deviation and vocal learning in a dataset of 164 mammals. Our work offers a testable framework for future empirical research linking allometric principles with the evolution of vocal learning.

Abstract

Communicating species identity is a key component of many animal signals. However, whether selection for species recognition systematically increases signal diversity during clade radiation remains debated. Here we show that in woodpecker drumming, a rhythmic signal used during mating and territorial defense, the amount of species identity information encoded remained stable during woodpeckers’ radiation. Acoustic analyses and evolutionary reconstructions show interchange among six main drumming types despite strong phylogenetic contingencies, suggesting evolutionary tinkering of drumming structure within a constrained acoustic space. Playback experiments and quantification of species discriminability demonstrate sufficient signal differentiation to support species recognition in local communities. Finally, we only find character displacement in the rare cases where sympatric species are also closely related. Overall, our results illustrate how historical contingencies and ecological interactions can promote conservatism in signals during a clade radiation without impairing the effectiveness of information transfer relevant to inter-specific discrimination.

Abstract

It is a common finding across languages that young children have problems in understanding patient-initial sentences. We used Tagalog, a verb-initial language with a reliable voice-marking system and highly frequent patient voice constructions, to test the predictions of several accounts that have been proposed to explain this difficulty: the frequency account, the Competition Model, and the incremental processing account. Study 1 presents an analysis of Tagalog child-directed speech, which showed that the dominant argument order is agent-before-patient and that morphosyntactic markers are highly valid cues to thematic role assignment. In Study 2, we used a combined self-paced listening and picture verification task to test how Tagalog-speaking adults and 5- and 7-year-old children process reversible transitive sentences. Results showed that adults performed well in all conditions, while children’s accuracy and listening times for the first noun phrase indicated more difficulty in interpreting patient-initial sentences in the agent voice compared to the patient voice. The patient voice advantage is partly explained by both the frequency account and incremental processing account.

Abstract

The constraints that govern acceptable phoneme combinations in speech perception and production have considerable plasticity. We addressed whether sleep influences the acquisition of new constraints and their integration into the speech-production system. Participants repeated sequences of syllables in which two phonemes were artificially restricted to syllable onset or syllable coda, depending on the vowel in that sequence. After 48 sequences, participants either had a 90-min nap or remained awake. Participants then repeated 96 sequences so implicit constraint learning could be examined, and then were tested for constraint generalization in a forced-choice task. The sleep group, but not the wake group, produced speech errors at test that were consistent with restrictions on the placement of phonemes in training. Furthermore, only the sleep group generalized their learning to new materials. Polysomnography data showed that implicit constraint learning was associated with slow-wave sleep. These results show that sleep facilitates the integration of new linguistic knowledge with existing production constraints. These data have relevance for systems-consolidation models of sleep.

Abstract

Processing of recursion has been proposed as the foundation of human linguistic ability. Yet this ability may be shared with other domains, such as the musical or rhythmic domain. Lindenmayer grammars (L-systems) have been proposed as a recursive grammar for use in artificial grammar experiments to test recursive processing abilities, and previous work had shown that participants are able to learn such a grammar using linguistic stimuli (syllables). In the present work, we used two experimental paradigms (a yes/no task and a two-alternative forced choice) to test whether adult participants are able to learn a recursive Lindenmayer grammar composed of drum sounds. After a brief exposure phase, we found that participants at the group level were sensitive to the exposure grammar and capable of distinguishing the grammatical and ungrammatical test strings above chance level in both tasks. While we found evidence of participants’ sensitivity to a very complex L-system grammar in a non-linguistic, potentially musical domain, the results were not robust. We discuss the discrepancy within our results and with the previous literature using L-systems in the linguistic domain. Furthermore, we propose directions for future music cognition research using L-system grammars.

Abstract

The way the human brain represents speech in memory is still unknown. An obvious characteristic of speech is its evolvement over time.
During speech processing, neural oscillations are modulated by the temporal properties of the acoustic speech signal, but also acquired
knowledge on the temporal structure of language influences speech perception-related brain activity. This suggests that speech could be
represented in the temporal domain, a form of representation that the brain also uses to encode autobiographic memories. Empirical
evidence for such a memory code is lacking. We investigated the nature of speech memory representations using direct cortical recordings
in the left perisylvian cortex during delayed sentence reproduction in female and male patients undergoing awake tumor surgery.
Our results reveal that the brain endogenously represents speech in the temporal domain. Temporal pattern similarity analyses revealed
that the phase of frontotemporal low-frequency oscillations, primarily in the beta range, represents sentence identity in working memory.
The positive relationship between beta power during working memory and task performance suggests that working memory
representations benefit from increased phase separation.

Abstract

Developmental dyslexia (DD) is one of the most prevalent learning disorders, with high impact on school and psychosocial development and high comorbidity with conditions like attention-deficit hyperactivity disorder (ADHD), depression, and anxiety. DD is characterized by deficits in different cognitive skills, including word reading, spelling, rapid naming, and phonology. To investigate the genetic basis of DD, we conducted a genome-wide association study (GWAS) of these skills within one of the largest studies available, including nine cohorts of reading-impaired and typically developing children of European ancestry (N = 2562–3468). We observed a genome-wide significant effect (p < 1 × 10−8) on rapid automatized naming of letters (RANlet) for variants on 18q12.2, within MIR924HG (micro-RNA 924 host gene; rs17663182 p = 4.73 × 10−9), and a suggestive association on 8q12.3 within NKAIN3 (encoding a cation transporter; rs16928927, p = 2.25 × 10−8). rs17663182 (18q12.2) also showed genome-wide significant multivariate associations with RAN measures (p = 1.15 × 10−8) and with all the cognitive traits tested (p = 3.07 × 10−8), suggesting (relational) pleiotropic effects of this variant. A polygenic risk score (PRS) analysis revealed significant genetic overlaps of some of the DD-related traits with educational attainment (EDUyears) and ADHD. Reading and spelling abilities were positively associated with EDUyears (p ~ [10−5–10−7]) and negatively associated with ADHD PRS (p ~ [10−8−10−17]). This corroborates a long-standing hypothesis on the partly shared genetic etiology of DD and ADHD, at the genome-wide level. Our findings suggest new candidate DD susceptibility genes and provide new insights into the genetics of dyslexia and its comorbities.

Abstract

Reading and language abilities are heritable traits that are likely to share some genetic influences with each other. To identify pleiotropic genetic variants affecting these traits, we first performed a Genome-wide Association Scan (GWAS) meta-analysis using three richly characterised datasets comprising individuals with histories of reading or language problems, and their siblings. GWAS was performed in a total of 1862 participants using the first principal component computed from several quantitative measures of reading- and language-related abilities, both before and after adjustment for performance IQ. We identified novel suggestive associations at the SNPs rs59197085 and rs5995177 (uncorrected p≈10−7 for each SNP), located respectively at the CCDC136/FLNC and RBFOX2 genes. Each of these SNPs then showed evidence for effects across multiple reading and language traits in univariate association testing against the individual traits. FLNC encodes a structural protein involved in cytoskeleton remodelling, while RBFOX2 is an important regulator of alternative splicing in neurons. The CCDC136/FLNC locus showed association with a comparable reading/language measure in an independent sample of 6434 participants from the general population, although involving distinct alleles of the associated SNP. Our datasets will form an important part of on-going international efforts to identify genes contributing to reading and language skills.

Abstract

Regional variation in African-American English (AAE) is especially salient to its speakers involved with hip-hop culture, as hip-hop assigns great importance to regional identity and regional accents are a key means of expressing regional identity. However, little is known about AAE regional variation regarding prosodic rhythm and melody. In hip-hop music, regional variation can also be observed, with different regions’ rap performances being characterized by distinct “flows” (i.e., rhythmic and melodic delivery), an observation which has not been quantitatively investigated yet. This study concerns regional variation in AAE speech and rap, specifically regarding the United States’ East and West Coasts. It investigates how East Coast and West Coast AAE prosody are distinct, how East Coast and West Coast rap flows differ, and whether the two domains follow a similar pattern: more rhythmic and melodic variation on the West Coast compared to the East Coast for both speech and rap. To this end, free speech and rap recordings of 16 prominent African-American members of the East Coast and West Coast hip-hop communities were phonetically analyzed regarding rhythm (e.g., syllable isochrony and musical timing) and melody (i.e., pitch fluctuation) using a combination of existing and novel methodological approaches. The results mostly confirm the hypotheses that East Coast AAE speech and rap are less rhythmically diverse and more monotone than West Coast AAE speech and rap, respectively. They also show that regional variation in AAE prosody and rap flows pattern in similar ways, suggesting a connection between rhythm and melody in language and music.

Abstract

Patterns formed by reaction-diffusion mechanisms are crucial for the development or sustenance of most organisms in nature. Patterns include dynamic waves, but are more often found as static distributions, such as animal skin patterns. Yet, a simplistic biological model system to reproduce and quantitatively investigate static reaction-diffusion patterns has been missing so far. Here, we demonstrate that the Escherichia coli MM system, known for its oscillatory behavior between the cell poles, is under certain conditions capable of transitioning to quasi-stationary protein distributions on membranes closely resembling Turing patterns. We systematically titrated both proteins, MinD and MinE, and found that removing all purification tags and linkers from the N-terminus of MinE was critical for static patterns to occur. At small bulk heights, dynamic patterns dominate, such as in rod-shaped microcompartments. We see implications of this work for studying pattern formation in general, but also for creating artificial gradients as downstream cues in synthetic biology applications.

Abstract

Interactive models of language production predict that it should be possible to observe long-distance interactions; effects that arise at one level of processing influence multiple subsequent stages of representation and processing. We examine the hypothesis that disruptions arising in nonform-based levels of planning—specifically, lexical selection—should modulate articulatory processing. A novel automatic phonetic analysis method was used to examine productions in a paradigm yielding both general disruptions to formulation processes and, more specifically, overt errors during lexical selection. This analysis method allowed us to examine articulatory disruptions at multiple levels of analysis, from whole words to individual segments. Baseline performance by young adults was contrasted with young speakers’ performance under time pressure (which previous work has argued increases interaction between planning and articulation) and performance by older adults (who may have difficulties inhibiting nontarget representations, leading to heightened interactive effects). The results revealed the presence of interactive effects. Our new analysis techniques revealed these effects were strongest in initial portions of responses, suggesting that speech is initiated as soon as the first segment has been planned. Interactive effects did not increase under response pressure, suggesting interaction between planning and articulation is relatively fixed. Unexpectedly, lexical selection disruptions appeared to yield some degree of facilitation in articulatory processing (possibly reflecting semantic facilitation of target retrieval) and older adults showed weaker, not stronger interactive effects (possibly reflecting weakened connections between lexical and form-level representations).

Abstract

Comparative thanatology encompasses the study of death-related responses in non-human animals and aspires to elucidate the evolutionary origins of human behavior in the context of death. Many reports have revealed that humans are not the only species affected by the death of group members. Non-human primates in particular show behaviors such as congregating around the deceased, carrying the corpse for prolonged periods of time (predominantly mothers carrying dead infants), and inspecting the corpse for signs of life. Here, we extend the focus on death-related responses in non-human animals by exploring whether chimpanzees are inclined to console the bereaved: the individual(s) most closely associated with the deceased. We report a case in which a chimpanzee (Pan troglodytes) mother experienced the loss of her fully developed infant (presumed stillborn). Using observational data to compare the group members’ behavior before and after the death, we found that a substantial number of group members selectively increased their affiliative expressions toward the bereaved mother. Moreover, on the day of the death, we observed heightened expressions of species-typical reassurance behaviors toward the bereaved mother. After ruling out several alternative explanations, we propose that many of the chimpanzees consoled the bereaved mother by means of affiliative and selective empathetic expressions.

Abstract

Chicks can convey information about their needs with calls. But it is still unknown if there are any universal need indicators in chick vocalizations. Previous studies have shown that in some species vocal activity and/or temporal-frequency variables of calls are related to the chick state, whereas other studies did not confirm it. Here, we tested experimentally whether vocal activity and temporal-frequency variables of calls change with cooling. We studied 10 human-raised
Siberian crane (Grus leucogeranus) chicks at 3–15 days of age. We found that the cooled chicks produced calls higher in fundamental
frequency and power variables, longer in duration and at a higher calling rate than in the control chicks. However, we did not find
significant changes in level of entropy and occurrence of non-linear phenomena in chick calls recorded during the experimental cooling. We suggest that the level of vocal activity is a universal indicator of need for warmth in precocial and semi-precocial birds (e.g. cranes), but not in altricial ones. We also assume that coding of needs via temporal-frequency variables of calls is typical in species whose adults could not confuse their chicks with other chicks. Siberian cranes stay on separate territories during their breeding season, so parents do not need to check individuality of their offspring in the home area. In this case, all call characteristics are available for other purposes and serve to communicate chicks’ vital needs.

Abstract

The present article examines the proposal that typology is a major factor guiding transfer selectivity in L3/Ln acquisition. We tested first exposure in L3/Ln using two artificial languages (ALs) lexically based in English and Spanish, focusing on gender agreement between determiners and nouns, and between nouns and adjectives. 50 L1 Spanish-L2 English speakers took part in the experiment. After receiving implicit training in one of the ALs (Mini-Spanish, N = 26; Mini-English, N = 24), gender violations elicited a fronto-lateral negativity in Mini-English in the earliest time window (200–500 ms), although this was not followed by any other differences in subsequent periods. This effect was highly localized, surfacing only in electrodes of the right-anterior region. In contrast, gender violations in Mini-Spanish elicited a broadly distributed positivity in the 300–600 ms time window. While we do not find typical indices of grammatical processing such as the P600 component, we believe that the between-groups differential appearance of the positivity for gender violations in the 300–600 ms time window reflects differential allocation of attentional resources as a function of the ALs’ lexical similarity to English or Spanish. We take these differences in attention to be precursors of the processes involved in transfer source selection in L3/Ln.

Abstract

Previous studies have described the existence of a phonotactic bias called the Labial–Coronal (LC) bias, corresponding to a tendency to produce more words beginning with a labial consonant followed by a coronal consonant (i.e. “bat”) than the opposite CL pattern (i.e. “tap”). This bias has initially been interpreted in terms of articulatory constraints of the human speech production system. However, more recently, it has been suggested that this presumably language-general LC bias in production might be accompanied by LC and CL biases in perception, acquired in infancy on the basis of the properties of the linguistic input. The present study investigates the origins of these perceptual biases, testing infants learning Japanese, a language that has been claimed to possess more CL than LC sequences, and comparing them with infants learning French, a language showing a clear LC bias in its lexicon. First, a corpus analysis of Japanese IDS and ADS revealed the existence of an overall LC bias, except for plosive sequences in ADS, which show a CL bias across counts. Second, speech preference experiments showed a perceptual preference for CL over LC plosive sequences (all recorded by a Japanese speaker) in 13- but not in 7- and 10-month-old Japanese-learning infants (Experiment 1), while revealing the emergence of an LC preference between 7 and 10 months in French-learning infants, using the exact same stimuli. These crosslinguistic behavioral differences, obtained with the same stimuli, thus reflect differences in processing in two populations of infants, which can be linked to differences in the properties of the lexicons of their respective native languages. These findings establish that the emergence of a CL/LC bias is related to exposure to a linguistic input.

Abstract

Humans associate abstract words with physical stimulus dimensions, such as linking upward locations with positive concepts (e.g., happy = up). These associations manifest both via subjective reports of associations and on objective performance metrics. Humans also report subjective associations between colours and abstract words (e.g., joy is linked to yellow). Here we tested whether such associations manifest on objective task performance, even when not task-relevant. Across three experiments, participants were presented with abstract words in physical colours that were either congruent with previously-reported subjective word-colour associations (e.g., victory in red and unhappy in blue), or were incongruent (e.g., victory in blue and unhappy in red). In Experiment 1, participants' task was to identify the valence of words. This congruency manipulation systematically affected objective task performance. In Experiment 2, participants completed two blocks, a valence-identification and a colour-identification task block. Both tasks produced congruency effects on performance, however, the results of the colour identification block could have reflected learning effects (i.e., associating the more common congruent colour with the word). This issue was rectified in Experiment 3, whereby participants completed the same two tasks as Experiment 2, but now matched congruent and incongruent pairs were used for both tasks. Again, both tasks produced reliable congruency effects. Item analyses in each experiment revealed that these effects demonstrated a degree of item specificity. Overall, there was clear evidence that at least some abstract word-colour pairings can systematically affect behaviour.

Abstract

Humans appear to rely on spatial mappings to represent and describe concepts. The conceptual cuing effect describes the tendency for participants to orient attention to a spatial location following the presentation of an unrelated cue word (e.g., orienting attention upward after reading the word sky). To date, such effects have predominately been explained within the embodied cognition framework, according to which people’s attention is oriented on the basis of prior experience (e.g., sky → up via perceptual simulation). However, this does not provide a compelling explanation for how abstract words have the same ability to orient attention. Why, for example, does dream also orient attention upward? We report on an experiment that investigated the role of language use (specifically, collocation between concept words and spatial words for up and down dimensions) and found that it predicted the cuing effect. The results suggest that language usage patterns may be instrumental in explaining conceptual cuing.

Abstract

Background: While persons with aphasia (PwA) are often dichotomised as fluent or nonfluent, agreement that fluency is not an all-or-nothing construct has led to the use of continuous variables as a way to quantify fluency, such as multi-dimensional rating scales, speech rate, and utterance length. Though these measures are often used in research, they provide little information about the underlying fluency deficit.
Aim: The aim of the study was to identify how well commonly used continuous measures of fluency capture variability in spontaneous speech variables at lexical, grammatical, and speech production levels. Methods & Procedures: Speech samples of 254 English-speaking PwA from the AphasiaBank database were analyzed to examine the distributions of four continuous measures of fluency: the WAB-R fluency scale, utterance length, retracing, and speech rate. Linear regression was used to identify spontaneous speech predictors contributing to each fluency outcome measure.
Outcomes & Results: All the outcome measures reflected the influence of multiple underlying dimensions, although the predictors varied. The WAB-R fluency scale, speech rate, and retracing were influenced by measures of grammatical competence, lexical retrieval, and speech production, whereas utterance length was influenced only by measures of grammatical competence and lexical retrieval. The strongest predictor of WAB-R fluency was aphasia severity, whereas the strongest predictor for all other fluency proxy measures was grammatical complexity.
Conclusions: Continuous measures allow a variety of ways to objectively quantify speech fluency; however, they reflect superficial manifestations of fluency that may be affected by multiple underlying deficits. Furthermore, the deficits underlying different measures vary, which may reduce the reliability of fluency diagnoses. Capturing these differences at the individual level is critical to accurate diagnosis and appropriately targeted therapy.

Abstract

Our aim in the present study is to measure neural correlates during spontaneous interactive sentence production. We present a novel approach using the word-by-word technique from improvisational theatre, in which two speakers jointly produce one sentence. This paradigm allows the assessment of behavioural aspects, such as turn-times, and electrophysiological responses, such as event-related-potentials (ERPs). Twenty-five participants constructed a cued but spontaneous four-word German sentence together with a confederate, taking turns for each word of the sentence. In 30% of the trials, the confederate uttered an unexpected gender-marked article. To complete the sentence in a meaningful way, the participant had to detect the violation and retrieve and utter a new fitting response. We found significant increases in response times after unexpected words and – despite allowing unscripted language production and naturally varying speech material – successfully detected significant N400 and P600 ERP effects for the unexpected word. The N400 EEG activity further significantly predicted the response time of the subsequent turn. Our results show that combining behavioural and neuroscientific measures of verbal interactions while retaining sufficient experimental control is possible, and that this combination provides promising insights into the mechanisms of spontaneous spoken dialogue.

Abstract

Our recent studies suggest that congenitally blind adults have severely impaired thresholds in an auditory spatial bisection task, pointing to the importance of vision in constructing complex auditory spatial maps (Gon etal., 2014). To explore strategies that may improve the auditory spatial sense in visually impaired people, we investigated the impact of tactile feedback on spatial auditory localization in 48 blindfolded sighted subjects. We measured auditory spatial bisection thresholds before and after training, either with tactile feedback, verbal feedback, or no feedback. Audio thresholds were first measured with a spatial bisection task: subjects judged whether the second sound of a three sound sequence was spatially closer to the first or the third sound. The tactile feedback group underwent two audio-tactile feedback sessions of 100 trials, where each auditory trial was followed by the same spatial sequence played on the subject's forearm; auditory spatial bisection thresholds were evaluated after each session. In the verbal feedback condition, the positions of the sounds were verbally reported to the subject after each feedback trial.The no feedback group did the same sequence of trials, with no feedback. Performance improved significantly only after audio-tactile feedback. The results suggest that direct tactile feedback interacts with the auditory spatial localization system, possibly by a process of cross-sensory recalibration. Control tests with the subject rotated suggested that this effect occurs only when the tactile and acoustic sequences are spatially congruent. Our results suggest that the tactile system can be used to recalibrate the auditory sense of space. These results encourage the possibility of designing rehabilitation programs to help blind persons establish a robust auditory sense of space, through training with the tactile modality.

Abstract

The aim of this study was to investigate whether early-English education benefits the perception
of English phonetic contrasts that are known to be perceptually confusable for Dutch
native speakers, comparing Dutch pupils who were enrolled in an early-English programme
at school from the age of four with pupils in a mainstream programme with English instruction
from the age of 11, and English-Dutch early bilingual children. Children were 4-5-yearolds
(start of primary school), 8-9-year-olds, or 11-12-year-olds (end of primary school).
Children were tested on four contrasts that varied in difficulty: /b/-/s/ (easy), /k/-/ɡ/ (intermediate),
/f/-/θ/ (difficult), /ε/-/æ/ (very difficult). Bilingual children outperformed the two other
groups on all contrasts except /b/-/s/. Early-English pupils did not outperform mainstream
pupils on any of the contrasts. This shows that early-English education as it is currently
implemented is not beneficial for pupils’ perception of non-native contrasts.

Abstract

In recent years, the influence of alpha (7–13 Hz) phase on visual processing has received a lot of attention. Magneto‐/encephalography (M/EEG) studies showed that alpha phase indexes visual excitability and task performance. Studies with transcranial alternating current stimulation (tACS) aim to modulate oscillations and causally impact task performance. Here, we applied right occipital tACS (O2 location) to assess the functional role of alpha phase in a series of experiments. We presented visual stimuli at different pre‐determined, experimentally controlled, phases of the entraining tACS signal, hypothesizing that this should result in an oscillatory pattern of visual performance in specifically left hemifield detection tasks. In experiment 1, we applied 10 Hz tACS and used separate psychophysical staircases for six equidistant tACS‐phase conditions, obtaining contrast thresholds for detection of visual gratings in left or right hemifield. In experiments 2 and 3, tACS was at EEG‐based individual peak alpha frequency. In experiment 2, we measured detection rates for gratings with (pseudo‐)fixed contrast. In experiment 3, participants detected brief luminance changes in a custom‐built LED device, at eight equidistant alpha phases. In none of the experiments did the primary outcome measure over phase conditions consistently reflect a one‐cycle sinusoid. However, post hoc analyses of reaction times (RT) suggested that tACS alpha phase did modulate RT for specifically left hemifield targets in both experiments 1 and 2 (not measured in experiment 3). This observation requires future confirmation, but is in line with the idea that alpha phase causally gates visual inputs through cortical excitability modulation.

Abstract

The cerebral cortex underlies our complex cognitive capabilities, yet little is known about the specific genetic loci that influence human cortical structure. To identify genetic variants that affect cortical structure, we conducted a genome-wide association meta-analysis of brain magnetic resonance imaging data from 51,665 individuals. We analyzed the surface area and average thickness of the whole cortex and 34 regions with known functional specializations. We identified 199 significant loci and found significant enrichment for loci influencing total surface area within regulatory elements that are active during prenatal cortical development, supporting the radial unit hypothesis. Loci that affect regional surface area cluster near genes in Wnt signaling pathways, which influence progenitor expansion and areal identity. Variation in cortical structure is genetically correlated with cognitive function, Parkinson’s disease, insomnia, depression, neuroticism, and attention deficit hyperactivity disorder.

Abstract

The involvement of neural motor and sensory systems in the processing of language has so far mainly been studied in native (L1) speakers. In an fMRI experiment, we investigated whether non-native (L2) semantic representations are rich enough to allow for activation in motor and somatosensory brain areas. German learners of Dutch and a control group of Dutch native speakers made lexical decisions about visually presented Dutch motor and non-motor verbs. Region-of-interest (ROI) and whole-brain analyses indicated that L2 speakers, like L1 speakers, showed significantly increased activation for simple motor compared to non-motor verbs in motor and somatosensory regions. This effect was not restricted to Dutch-German cognate verbs, but was also present for non-cognate verbs. These results indicate that L2 semantic representations are rich enough for motor-related activations to develop in motor and somatosensory areas.

Abstract

In this functional magnetic resonance imaging (fMRI) long-lag priming study, we investigated the processing of Dutch semantically transparent, derived prefix verbs. In such words, the meaning of the word as a whole can be deduced from the meanings of its parts, e.g., wegleggen “put aside.” Many behavioral and some fMRI studies suggest that native (L1) speakers decompose transparent derived words. The brain region usually implicated in morphological decomposition is the left inferior frontal gyrus (LIFG). In non-native (L2) speakers, the processing of transparent derived words has hardly been investigated, especially in fMRI studies, and results are contradictory: some studies find more reliance on holistic (i.e., non-decompositional) processing by L2 speakers; some find no difference between L1 and L2 speakers. In this study, we wanted to find out whether Dutch transparent derived prefix verbs are decomposed or processed holistically by German L2 speakers of Dutch. Half of the derived verbs (e.g., omvallen “fall down”) were preceded by their stem (e.g., vallen “fall”) with a lag of 4–6 words (“primed”); the other half (e.g., inslapen “fall asleep”) were not (“unprimed”). L1 and L2 speakers of Dutch made lexical decisions on these visually presented verbs. Both region of interest analyses and whole-brain analyses showed that there was a significant repetition suppression effect for primed compared to unprimed derived verbs in the LIFG. This was true both for the analyses over L2 speakers only and for the analyses over the two language groups together. The latter did not reveal any interaction with language group (L1 vs. L2) in the LIFG. Thus, L2 speakers show a clear priming effect in the LIFG, an area that has been associated with morphological decomposition. Our findings are consistent with the idea that L2 speakers engage in decomposition of transparent derived verbs rather than processing them holistically

Abstract

This study examined electrophysiological correlates of sentence comprehension of native-accented and foreign-accented
speech in a second language (L2), for sentences produced in a foreign accent different from that associated with the listeners’
L1. Bilingual speaker-listeners process different accents in their L2 conversations, but the effects on real-time L2 sentence
comprehension are unknown. Dutch–English bilinguals listened to native American-English accented sentences and foreign
(and for them unfamiliarly-accented) Chinese-English accented sentences while EEG was recorded. Behavioral sentence
comprehension was highly accurate for both native-accented and foreign-accented sentences. ERPs showed different patterns
for L2 grammar and semantic processing of native- and foreign-accented speech. For grammar, only native-accented speech
elicited an Nref. For semantics, both native- and foreign-accented speech elicited an N400 effect, but with a delayed onset
across both accent conditions. These findings suggest that the way listeners comprehend native- and foreign-accented
sentences in their L2 depends on their familiarity with the accent.

Abstract

Autism spectrum disorder (ASD) is a highly heritable and heterogeneous group of neurodevelopmental phenotypes diagnosed in more than 1% of children. Common genetic variants contribute substantially to ASD susceptibility, but to date no individual variants have been robustly associated with ASD. With a marked sample-size increase from a unique Danish population resource, we report a genome-wide association meta-analysis of 18,381 individuals with ASD and 27,969 controls that identified five genome-wide-significant loci. Leveraging GWAS results from three phenotypes with significantly overlapping genetic architectures (schizophrenia, major depression, and educational attainment), we identified seven additional loci shared with other traits at equally strict significance levels. Dissecting the polygenic architecture, we found both quantitative and qualitative polygenic heterogeneity across ASD subtypes. These results highlight biological insights, particularly relating to neuronal function and corticogenesis, and establish that GWAS performed at scale will be much more productive in the near term in ASD.

Abstract

The left and right sides of the human brain are specialized for different kinds of information processing, and much of our cognition is lateralized to an extent towards one side or the other. Handedness is a reflection of nervous system lateralization. Roughly ten percent of people are mixed- or left-handed, and they show an elevated rate of reductions or reversals of some cerebral functional asymmetries compared to right-handers. Brain anatomical correlates of left-handedness have also been suggested. However, the relationships of left-handedness to brain structure and function remain far from clear. We carried out a comprehensive analysis of cortical surface area differences between 106 left-handed subjects and 1960 right-handed subjects, measured using an automated method of regional parcellation (FreeSurfer, Destrieux atlas). This is the largest study sample that has so far been used in relation to this issue. No individual cortical region showed an association with left-handedness that survived statistical correction for multiple testing, although there was a nominally significant association with the surface area of a previously implicated region: the left precentral sulcus. Identifying brain structural correlates of handedness may prove useful for genetic studies of cerebral asymmetries, as well as providing new avenues for the study of relations between handedness, cerebral lateralization and cognition.

Abstract

Functional and anatomical asymmetries are prevalent features of the human brain, linked to gender, handedness, and cognition. However, little is known about the neurodevelopmental processes involved. In zebrafish, asymmetries arise in the diencephalon before extending within the central nervous system. We aimed to identify genes involved in the development of subtle, left-right volumetric asymmetries of human subcortical structures using large datasets. We first tested the feasibility of measuring left-right volume differences in such large-scale samples, as assessed by two automated methods of subcortical segmentation (FSL|FIRST and FreeSurfer), using data from 235 subjects who had undergone MRI twice. We tested the agreement between the first and second scan, and the agreement between the segmentation methods, for measures of bilateral volumes of six subcortical structures and the hippocampus, and their volumetric asymmetries. We also tested whether there were biases introduced by left-right differences in the regional atlases used by the methods, by analyzing left-right flipped images. While many bilateral volumes were measured well (scan-rescan r = 0.6-0.8), most asymmetries, with the exception of the caudate nucleus, showed lower repeatabilites. We meta-analyzed genome-wide association scan results for caudate nucleus asymmetry in a combined sample of 3,028 adult subjects but did not detect associations at genome-wide significance (P < 5 × 10-8). There was no enrichment of genetic association in genes involved in left-right patterning of the viscera. Our results provide important information for researchers who are currently aiming to carry out large-scale genome-wide studies of subcortical and hippocampal volumes, and their asymmetries

Abstract

A large body of evidence has shown that visual context information can rapidly modulate language comprehension for concrete sentences and when it is mediated by a referential or a lexical-semantic link. What has not yet been examined is whether visual context can also modulate comprehension of abstract sentences incrementally when it is neither referenced by, nor lexically associated with, the sentence. Three eye-tracking reading experiments examined the effects of spatial distance between words (Experiment 1) and objects (Experiment 2 and 3) on participants’ reading times for sentences that convey similarity or difference between two abstract nouns (e.g., ‘Peace and war are certainly different...’). Before reading the sentence, participants inspected a visual context with two playing cards that moved either far apart or close together. In Experiment 1, the cards turned and showed the first two nouns of the sentence (e.g., ‘peace’, ‘war’). In Experiments 2 and 3, they turned but remained blank. Participants’ reading times at the adjective (Experiment 1: first-pass reading time; Experiment 2: total times) and at the second noun phrase (Experiment 3: first-pass times) were faster for sentences that expressed similarity when the preceding words/objects were close together (vs. far apart) and for sentences that expressed dissimilarity when the preceding words/objects were far apart (vs. close together). Thus, spatial distance between words or entirely unrelated objects can rapidly and incrementally modulate the semantic interpretation of abstract sentences.

Abstract

Partisan gerrymandering poses a threat to democracy. Moreover, the complexity of the districting task may exceed human capacities. One potential solution is using computational models to automate the districting process by optimizing objective and open criteria, such as how spatially compact districts are. We formulated one such model that minimised pairwise distance between voters within a district. Using US Census Bureau data, we confirmed our prediction that the difference in compactness between the computed and actual districts would be greatest for states that are large and, therefore, difficult for humans to properly district given their limited capacities. The computed solutions highlighted differences in how humans and machines solve this task with machine solutions more fully optimised and displaying emergent properties not evident in human solutions. These results suggest a division of labour in which humans debate and formulate districting criteria whereas machines optimise the criteria to draw the district boundaries. We discuss how criteria can be expanded beyond notions of compactness to include other factors, such as respecting municipal boundaries, historic communities, and relevant legislation.

Abstract

A key strength of connectionist modelling is its ability to simulate both intact cognition and the behavioural effects of neural damage. We survey the literature, showing that models have been damaged in a variety of ways, e.g. by removing connections, by adding noise to connection weights, by scaling weights, by removing units and by adding noise to unit activations. While these different implementations of damage have often been assumed to be behaviourally equivalent, some theorists have made aetiological claims that rest on nonequivalence. They suggest that related deficits with different aetiologies might be accounted for by different forms of damage within a single model. We present two case studies that explore the effects of different forms of damage in two influential connectionist models, each of which has been applied to explain neuropsychological deficits. Our results indicate that the effect of simulated damage can indeed be sensitive to the way in which damage is implemented, particularly when the environment comprises subsets of items that differ in their statistical properties, but such effects are sensitive to relatively subtle aspects of the model’s training environment. We argue that, as a consequence, substantial methodological care is required if aetiological claims about simulated neural damage are to be justified, and conclude more generally that implementation assumptions, including those concerning simulated damage, must be fully explored when evaluating models of neurological deficits, both to avoid over-extending the explanatory power of specific implementations and to ensure that reported results are replicable.

Abstract

Since listeners usually look at the speaker's face, gestural information has to be absorbed through peripheral visual perception. In the literature, it has been suggested that listeners look at gestures under certain circumstances: 1) when the articulation of the gesture is peripheral; 2) when the speech channel is insufficient for comprehension; and 3) when the speaker him- or herself indicates that the gesture is worthy of attention. The research here reported employs eye tracking techniques to study the perception of gestures in face-to-face interaction. The improved control over the listener's visual channel allows us to test the validity of the above claims. We present preliminary findings substantiating claims 1 and 3, and relate them to theoretical proposals in the literature and to the issue of how visual and cognitive attention are related.

Abstract

One of the features that distinguishes modern humans from our extinct relatives
and ancestors is a globular shape of the braincase [1-4]. As the endocranium
closely mirrors the outer shape of the brain, these differences might reflect
altered neural architecture [4,5]. However, in the absence of fossil brain tissue the
underlying neuroanatomical changes as well as their genetic bases remain
elusive. To better understand the biological foundations of modern human
endocranial shape, we turn to our closest extinct relatives, the Neandertals.
Interbreeding between modern humans and Neandertals has resulted in
introgressed fragments of Neandertal DNA in the genomes of present-day non-
Africans [6,7]. Based on shape analyses of fossil skull endocasts, we derive a
measure of endocranial globularity from structural magnetic resonance imaging
(MRI) scans of thousands of modern humans, and study the effects of
introgressed fragments of Neandertal DNA on this phenotype. We find that
Neandertal alleles on chromosomes 1 and 18 are associated with reduced
endocranial globularity. These alleles influence expression of two nearby genes,
UBR4 and PHLPP1, which are involved in neurogenesis and myelination,
respectively. Our findings show how integration of fossil skull data with archaic
genomics and neuroimaging can suggest developmental mechanisms that may
contribute to the unique modern human endocranial shape.

Abstract

What is the long‐term trajectory of semantic memory deficits in patients who have suffered structural brain damage? Memory is, per definition, a changing faculty. The traditional view is that after an initial recovery period, the mature human brain has little capacity to repair or reorganize. More recently, it has been suggested that the central nervous system may be more plastic with the ability to change in neural structure, connectivity, and function. The latter observations are, however, largely based on normal learning in healthy subjects. Here, we report a patient who suffered bilateral ventro‐medial damage after presumed herpes encephalitis in 1971. He was seen regularly in the eighties, and we recently had the opportunity to re‐assess his semantic memory deficits. On semantic category fluency, he showed a very clear category‐specific deficit performing better that control data on non‐living categories and significantly worse on living items. Recent testing showed that his impairments have remained unchanged for more than 40 years. We suggest cautiousness when extrapolating the concept of brain plasticity, as observed during normal learning, to plasticity in the context of structural brain damage.

Abstract

The central issue of this study concerns the claim that the processing of gender agreement in online sentence comprehension is a syntactic rather than a conceptual/semantic process. This claim was tested for the grammatical gender agreement in Dutch between the definite article and the noun. Subjects read sentences in which the definite article and the noun had the same gender and sentences in which the gender agreement was violated, While subjects read these sentences, their electrophysiological activity was recorded via electrodes placed on the scalp. Earlier research has shown that semantic and syntactic processing events manifest themselves in different event-related brain potential (ERP) effects. Semantic integration modulates the amplitude of the so-called N400.The P600/SPS is an ERP effect that is more sensitive to syntactic processes. The violation of grammatical gender agreement was found to result in a P600/SPS. For violations in sentence-final position, an additional increase of the N400 amplitude was observed. This N400 effect is interpreted as resulting from the consequence of a syntactic violation for the sentence-final wrap-up. The overall pattern of results supports the claim that the on-line processing of gender agreement information is not a content driven but a syntactic-form driven process.

Abstract

Lexical-semantic processing impairments in aphasic patients with left hemisphere lesions and non-aphasic patients with right hemisphere lesions were investigated by recording event-related brain potentials (ERPs) while subjects listened to auditorily presented word pairs. The word pairs consisted of unrelated words, or words that were related in meaning. The related words were either associatively related, e.g. 'bread-butter', or were members of the same semantic category without being associatively related, e.g. 'churchvilla '. The latter relationships are assumed to be more distant than the former ones. The most relevant ERP component in this study is the N400. In elderly control subjects, the N400 amplitude to associatively and semantically related word targets is reduced relative to the N400 elicited by unrelated targets. Compared with this normal N400 effect, the different patient groups showed the following pattern of results: aphasic patients with only minor comprehension deficits (high comprehenders) showed N400 effects of a similar size as the control subjects. In aphasic patients with more severe comprehension deficits (low comprehenders) a clear reduction in the N400 effects was obtained, both for the associative and the semantic word pairs. The patients with right hemisphere lesions showed a normal N400 effect for the associatively related targets, but a trend towards a reduced N400 effect for the semantically related word pairs. A dissociation between the N400 results in the word pair paradigm and P300 results in a classical tone oddball task indicated that the N400 effects were not an aspecific consequence of brain lesion, but were related to the nature of the language comprehension impairment. The conclusions drawn from the ERP results are that comprehension deficits in the aphasic patients are due to an impairment in integrating individual word meanings into an overall meaning representation. Right hemisphere patients are more specifically impaired in the processing of semantically more distant relationships, suggesting the involvement of the right hemisphere in semantically coarse coding.

Abstract

Current views on the neurobiological underpinnings of language are discussed that deviate in a number of ways from the classical Wernicke–Lichtheim–Geschwind model. More areas than Broca's and Wernicke's region are involved in language. Moreover, a division along the axis of language production and language comprehension does not seem to be warranted. Instead, for central aspects of language processing neural infrastructure is shared between production and comprehension. Three different accounts of the role of Broca's area in language are discussed. Arguments are presented in favor of a dynamic network view, in which the functionality of a region is co-determined by the network of regions in which it is embedded at particular moments in time. Finally, core regions of language processing need to interact with other networks (e.g. the attentional networks and the ToM network) to establish full functionality of language and communication.

Abstract

In this contribution, the following four questions are discussed: (i) where is meaning?; (ii) what is meaning?; (iii) what is the meaning of mechanism?; (iv) what are the mechanisms of meaning? I will argue that meanings are in the head. Meanings have multiple facets, but minimally one needs to make a distinction between single word meanings (lexical meaning) and the meanings of multi-word utterances. The latter ones cannot be retrieved from memory, but need to be constructed on the fly. A mechanistic account of the meaning-making mind requires an analysis at both a functional and a neural level, the reason being that these levels are causally interdependent. I will show that an analysis exclusively focusing on patterns of brain activation lacks explanatory power. Finally, I shall present an initial sketch of how the dynamic interaction between temporo-parietal areas and inferior frontal cortex might instantiate the interpretation of linguistic utterances in the context of a multimodal setting and ongoing discourse information.

Abstract

Silent reading and reading aloud of German words and pseudowords were used in a PET study using (15O)butanol to examine the neural correlates of reading and of the phonological conversion of legal letter strings, with or without meaning.
The results of 11 healthy, right-handed volunteers in the age range of 25 to 30 years showed activation of the lingual gyri during silent reading in comparison with viewing a fixation cross. Comparisons between the reading of words and pseudowords suggest the involvement of the middle temporal gyri in retrieving both the phonological and semantic code for words. The reading of pseudowords activates the left inferior frontal gyrus, including the ventral part of Broca’s area, to a larger extent than the reading of words. This suggests that this area might be involved in the sublexical conversion of orthographic input strings into phonological output codes. (Pre)motor areas were found to be activated during both silent reading and reading aloud. On the basis of the obtained activation patterns, it is hypothesized that the articulation of high-frequency syllables requires the retrieval of their concomitant articulatory gestures from the SMA and that the articulation of lowfrequency syllables recruits the left medial premotor cortex.

Abstract

In this Review, I propose a multiple-network view for the neurobiological basis of distinctly human language skills. A much more complex picture of interacting brain areas emerges than in the classical neurobiological model of language. This is because using language is more than single-word processing, and much goes on beyond the information given in the acoustic or orthographic tokens that enter primary sensory cortices. This requires the involvement of multiple networks with functionally nonoverlapping contributions

Abstract

A hallmark of human language is that we combine lexical building blocks retrieved from memory in endless new ways. This combinatorial aspect of language is referred to as unification. Here we focus on the neurobiological infrastructure for syntactic and semantic unification. Unification is characterized by a high-speed temporal profile including both prediction and integration of retrieved lexical elements. A meta-analysis of numerous neuroimaging studies reveals a clear dorsal/ventral gradient in both left inferior frontal cortex and left posterior temporal cortex, with dorsal foci for syntactic processing and ventral foci for semantic processing. In addition to core areas for unification, further networks need to be recruited to realize language-driven communication to its full extent. One example is the theory of mind network, which allows listeners and readers to infer the intended message (speaker meaning) from the coded meaning of the linguistic utterance. This indicates that sensorimotor simulation cannot handle all of language processing.

Abstract

Children’s songs are a great source for linguistic learning. Here we explore whether children can acquire novel words in a second language by playing a game featuring children’s songs in a playhouse. The playhouse is designed by the Noplica foundation (www.noplica.nl) to advance language learning through unsupervised play. We present data from three experiments that serve to scientifically proof the functionality of one game of the playhouse: the Energy Center. For this game, children move three hand-bikes mounted on a panel within the playhouse. Once the children cycle, a song starts playing that is accompanied by musical instruments. In our experiments, children executed a picture-selection task to evaluate whether they acquired new vocabulary from the songs presented during the game. Two of our experiments were run in the field, one at a Dutch and one at an Indian pre-school. The third experiment features data from a more controlled laboratory setting. Our results partly confirm that the Energy Center is a successful means to support vocabulary acquisition in a second language. More research with larger sample sizes and longer access to the Energy Center is needed to evaluate the overall functionality of the game. Based on informal observations at our test sites, however, we are certain that children do pick up linguistic content from the songs during play, as many of the children repeat words and phrases from the songs they heard. We will pick up upon these promising observations during future studies.

Abstract

Infants exploit acoustic boundaries to perceptually organize phrases in speech. This prosodic parsing ability is well‐attested and is a cornerstone to the development of speech perception and grammar. However, infants also receive linguistic input in child songs. This study provides evidence that infants parse songs into meaningful phrasal units and replicates previous research for speech. Six‐month‐old Dutch infants (n = 80) were tested in the song or speech modality in the head‐turn preference procedure. First, infants were familiarized to two versions of the same word sequence: One version represented a well‐formed unit, and the other contained a phrase boundary halfway through. At test, infants were presented two passages, each containing one version of the familiarized sequence. The results for speech replicated the previously observed preference for the passage containing the well‐formed sequence, but only in a more fine‐grained analysis. The preference for well‐formed phrases was also observed in the song modality, indicating that infants recognize phrase structure in song. There were acoustic differences between stimuli of the current and previous studies, suggesting that infants are flexible in their processing of boundary cues while also providing a possible explanation for differences in effect sizes.

Abstract

In the recent past the work on large-scale linguistic distributions across the globe has intensified considerably. Work on macro-areal relationships in Africa (Güldemann, 2010) suggests that the shape of convergence areas may be determined by climatic factors and geophysical features such as mountains, water bodies, coastlines, etc. Worldwide data is now available for geophysical features as well as linguistic features, including numeral systems and basic constituent order. We explore the possibility that the shape of areal aggregations of individual features in these two linguistic domains correlates with Köppen-Geiger climate zones. Furthermore, we test the hypothesis that the shape of such areal feature aggregations is determined by the contour of adjacent geophysical features like mountain ranges or coastlines. In these first basic tests, we do not find clear evidence that either Köppen-Geiger climate zones or the contours of geophysical features are major predictors for the linguistic data at hand

Abstract

While the notion of the ‘area’ or ‘Sprachbund’ has a long history in linguistics, with geographically-defined regions frequently cited as a useful means to explain typological distributions, the problem of delimiting areas has not been well addressed. Lists of general-purpose, largely independent ‘macro-areas’ (typically continent size) have been proposed as a step to rule out contact as an explanation for various large-scale linguistic phenomena. This squib points out some problems in some of the currently widely-used predetermined areas, those found in the World Atlas of Language Structures (Haspelmath et al., 2005). Instead, we propose a principled division of the world’s landmasses into six macro-areas that arguably have better geographical independence properties

Abstract

Speech production studies have shown that phonological unit initially used to fill the metrical frame during phonological encoding is language specific, that is, a phoneme for English and Dutch, an atonal syllable for Mandarin Chinese, and a mora for Japanese. However, only a few studies chronometrically investigated speech production in Korean, and they obtained mixed results. Korean is particularly interesting as there might be both phonemic and syllabic influences during phonological encoding. The purpose of this study is to further examine the initial phonological preparation unit in Korean, employing a masked priming task (Experiment 1) and a phonological Stroop task (Experiment 2). The results showed that significant onset (and onset-plus, that is, consonant-vowel [CV]) effects were found in both experiments, but there was no compelling evidence for a prominent role for the syllable. When the prime words were presented in three different forms related to the targets, namely, without any change, with re-syllabified codas, and with nasalised codas, there were no significant differences in facilitation among the three forms. Alternatively, it is also possible that participants may not have had sufficient time to process the primes up to the point that re-syllabification or nasalisation could have been carried out. In addition, the results of a Stroop task demonstrated that the onset phoneme effect was not driven by any orthographic influence. These findings suggest that the onset segment and not the syllable is the initial (or proximate) phonological unit used in the segment-to-frame encoding process during speech planning in Korean.

Abstract

Feedback has been shown to be effective in shifting attention across perceptual cues to a phonological contrast in speech perception (Francis, Baldwin & Nusbaum, 2000). However, the learning mechanisms behind this process remain obscure. We compare the predictions of supervised error-driven learning (Rescorla & Wagner, 1972) and reinforcement learning (Sutton & Barto, 1998) using computational simulations. Supervised learning predicts downweighting of an informative cue when the learner receives evidence that it is no longer informative. In contrast, reinforcement learning suggests that a reduction in cue weight requires positive evidence for the informativeness of an alternative cue. Experimental evidence supports the latter prediction, implicating reinforcement learning as the mechanism behind the effect of feedback on cue weighting in speech perception. Native English listeners were exposed to either bimodal or unimodal VOT distributions spanning the unaspirated/aspirated boundary (bear/pear). VOT is the primary cue to initial stop voicing in English. However, lexical feedback in training indicated that VOT was no longer predictive of voicing. Reduction in the weight of VOT was observed only when participants could use an alternative cue, F0, to predict voicing. Frequency distributions had no effect on learning. Overall, the results suggest that attention shifting in learning the phonetic cues to phonological categories is accomplished using simple reinforcement learning principles that also guide the choice of actions in other domains.

Abstract

Previous studies have linked the low expression variant of a variable number of tandem repeat polymorphism in the monoamine oxidase A gene (MAOA‐L) to the risk for impulsivity and aggression, brain developmental abnormalities, altered cortico‐limbic circuit function, and an exaggerated neural serotonergic tone. However, the neurobiological effects of this variant on human brain network architecture are incompletely understood. We studied healthy individuals and used multimodal neuroimaging (sample size range: 219–284 across modalities) and network‐based statistics (NBS) to probe the specificity of MAOA‐L‐related connectomic alterations to cortical‐limbic circuits and the emotion processing domain. We assessed the spatial distribution of affected links across several neuroimaging tasks and data modalities to identify potential alterations in network architecture. Our results revealed a distributed network of node links with a significantly increased connectivity in MAOA‐L carriers compared to the carriers of the high expression (H) variant. The hyperconnectivity phenotype primarily consisted of between‐lobe (“anisocoupled”) network links and showed a pronounced involvement of frontal‐temporal connections. Hyperconnectivity was observed across functional magnetic resonance imaging (fMRI) of implicit emotion processing (pFWE = .037), resting‐state fMRI (pFWE = .022), and diffusion tensor imaging (pFWE = .044) data, while no effects were seen in fMRI data of another cognitive domain, that is, spatial working memory (pFWE = .540). These observations are in line with prior research on the MAOA‐L variant and complement these existing data by novel insights into the specificity and spatial distribution of the neurogenetic effects. Our work highlights the value of multimodal network connectomic approaches for imaging genetics.

Abstract

All primates learn things from conspecifics socially, but it is not clear whether they conform to the behavior of these conspecifics—if conformity is defined as overriding individually acquired behavioral tendencies in order to copy peers’ behavior. In the current study, chimpanzees, orangutans, and 2-year-old human children individually acquired a problem-solving strategy. They then watched several conspecific peers demonstrate an alternative strategy. The children switched to this new, socially demonstrated strategy in roughly half of all instances, whereas the other two great-ape species almost never adjusted their behavior to the majority’s. In a follow-up study, children switched much more when the peer demonstrators were still present than when they were absent, which suggests that their conformity arose at least in part from social motivations. These results demonstrate an important difference between the social learning of humans and great apes, a difference that might help to account for differences in human and nonhuman cultures

Abstract

Preregistration, the act of specifying a research plan in advance, is becoming more common in scientific research. Infant researchers contend with unique problems that might make preregistration particularly challenging. Infants are a hard‐to‐reach population, usually yielding small sample sizes, they can only complete a limited number of trials, and they can be excluded based on hard‐to‐predict complications (e.g., parental interference, fussiness). In addition, as effects themselves potentially change with age and population, it is hard to calculate an a priori effect size. At the same time, these very factors make preregistration in infant studies a valuable tool. A priori examination of the planned study, including the hypotheses, sample size, and resulting statistical power, increases the credibility of single studies and adds value to the field. Preregistration might also improve explicit decision making to create better studies. We present an in‐depth discussion of the issues uniquely relevant to infant researchers, and ways to contend with them in preregistration and study planning. We provide recommendations to researchers interested in following current best practices.

Abstract

Cranial growth and development is a complex process which affects the closely related traits of head circumference (HC) and intracranial volume (ICV). The underlying genetic influences affecting these traits during the transition from childhood to adulthood are little understood, but might include both age-specific genetic influences and low-frequency genetic variation. To understand these influences, we model the developmental genetic architecture of HC, showing this is genetically stable and correlated with genetic determinants of ICV. Investigating up to 46,000 children and adults of European descent, we identify association with final HC and/or final ICV+HC at 9 novel common and low-frequency loci, illustrating that genetic variation from a wide allele frequency spectrum contributes to cranial growth. The largest effects are reported for low-frequency variants within TP53, with 0.5 cm wider heads in increaser-allele carriers versus non-carriers during mid-childhood, suggesting a previously unrecognized role of TP53 transcripts in human cranial development.

Abstract

The present article on executive control addresses the issue of the locus of the Stroop effect by examining neurophysiological components marking conflict monitoring, interference suppression, and conflict resolution. Our goal was to provide an overview of a series of determining neurophysiological findings including neural source reconstruction data on distinct executive control processes and sub-processes involved in the Stroop task. Consistently, a fronto-central N2 component is found to reflect conflict monitoring processes, with its main neural generator being the anterior cingulate cortex (ACC). Then, for cognitive control tasks that involve a linguistic component like the Stroop task, the N2 is followed by a centro-posterior N400 and subsequently a late sustained potential (LSP). The N400 is mainly generated by the ACC and the prefrontal cortex (PFC) and is thought to reflect interference suppression, whereas the LSP plausibly reflects conflict resolution processes. The present overview shows that ERP constitute a reliable methodological tool for tracing with precision the time course of different executive processes and sub-processes involved in experimental tasks involving a cognitive conflict. Future research should shed light on the fine-grained mechanisms of control respectively involved in linguistic and non-linguistic tasks.

Abstract

Speakers and listeners usually interact in larger discourses than single words or even single sentences. The goal of the present study was to identify the neural bases reflecting how the mental representation of the situation denoted in a multi-sentence discourse (situation model) is constructed and shared between speakers and listeners. An fMRI study using a variant of the ambiguous text paradigm was designed. Speakers (n=15) produced ambiguous texts in the scanner and listeners (n=27) subsequently listened to these texts in different states of ambiguity: preceded by a highly informative, intermediately informative or no title at all. Conventional BOLD activation analyses in listeners, as well as inter-subject correlation analyses between the speakers’ and the listeners’ hemodynamic time courses were performed. Critically, only the processing of disambiguated, coherent discourse with an intelligible situation model representation involved (shared) activation in bilateral lateral parietal and medial prefrontal regions. This shared spatiotemporal pattern of brain activation between the speaker and the listener suggests that the process of memory retrieval in medial prefrontal regions and the binding of retrieved information in the lateral parietal cortex constitutes a core mechanism underlying the communication of complex conceptual representations.

Abstract

Visual context facilitates perception, but how this is neurally implemented remains unclear. One example of contextual facilitation is found in reading, where letters are more easily identified when embedded in a word. Bottom-up models explain this word advantage as a post-perceptual decision bias, while top-down models propose that word contexts enhance perception itself. Here, we arbitrate between these accounts by presenting words and nonwords and probing the representational fidelity of individual letters using functional magnetic resonance imaging. In line with top-down models, we find that word contexts enhance letter representations in early visual cortex. Moreover, we observe increased coupling between letter information in visual cortex and brain activity in key areas of the reading network, suggesting these areas may be the source of the enhancement. Our results provide evidence for top-down representational enhancement in word recognition, demonstrating that word contexts can modulate perceptual processing already at the earliest visual regions.

Abstract

Timing is an essential parameter influencing many behaviours. A previous study demonstrated a high sensitivity of a phocid, the harbour seal (Phoca vitulina), in discriminating time intervals. In the present study, we compared the harbour seal’s timing abilities with the timing abilities of an otariid, the South African fur seal (Arctocephalus pusillus pusillus). This comparison seemed essential as phocids and otariids differ in many respects and might, thus, also differ regarding their timing abilities. We determined time difference thresholds for sub- and suprasecond time intervals marked by a white circle on a black background displayed for a specific time interval on a monitor using a staircase method. Contrary to our expectation, the timing abilities of the fur seal and the harbour seal were comparable. Over a broad range of time intervals, 0.8–7 s in the fur seal and 0.8–30 s in the harbour seal, the difference thresholds followed Weber’s law. In this range, both animals could discriminate time intervals differing only by 12 % and 14 % on average. Timing might, thus be a fundamental cue for pinnipeds in general to be used in various contexts, thereby complementing information provided by classical sensory systems. Future studies will help to clarify if timing is indeed involved in foraging decisions or the estimation of travel speed or distance.

Abstract

Polymorphisms in the apolipoprotein E (APOE) gene have been associated with individual differences in cognition, brain structure and brain function. For example, the ε4 allele has been associated with cognitive and brain impairment in old age and increased risk of dementia, while the ε2 allele has been claimed to be neuroprotective. According to the ‘antagonistic pleiotropy’ hypothesis, these polymorphisms have different effects across the lifespan, with ε4, for example, postulated to confer benefits on cognitive and brain functions earlier in life. In this stage 2 of the Registered Report – https://osf.io/bufc4, we report the results from the cognitive and brain measures in the Cambridge Centre for Ageing and Neuroscience cohort (www.cam-can.org). We investigated the antagonistic pleiotropy hypothesis by testing for allele-by-age interactions in approximately 600 people across the adult lifespan (18–88 years), on six outcome variables related to cognition, brain structure and brain function (namely, fluid intelligence, verbal memory, hippocampal grey-matter volume, mean diffusion within white matter and resting-state connectivity measured by both functional magnetic resonance imaging and magnetoencephalography). We found no evidence to support the antagonistic pleiotropy hypothesis. Indeed, Bayes factors supported the null hypothesis in all cases, except for the (linear) interaction between age and possession of the ε4 allele on fluid intelligence, for which the evidence for faster decline in older ages was ambiguous. Overall, these pre-registered analyses question the antagonistic pleiotropy of APOE polymorphisms, at least in healthy adults.

Abstract

Bilingual listeners comprehend speech-in-noise better in their native than non-native language. This native-language benefit is thought to arise from greater use of top-down linguistic information to assist degraded speech comprehension. Using functional magnetic resonance imaging, we recently showed that left angular gyrus activation is modulated when semantic context is used to assist native language speech-in-noise comprehension (Golestani, Hervais-Adelman, Obleser, & Scott, 2013). Here, we extend the previous work, by reanalyzing the previous data alongside the results obtained in the non-native language of the same late bilingual participants. We found a behavioral benefit of semantic context in processing speech-in-noise in the native language only, and the imaging results also revealed a native language context effect in the left angular gyrus. We also find a complementary role of lower-level auditory regions during stimulus-driven processing. Our findings help to elucidate the neural basis of the established native language behavioral benefit of speech-in-noise processing. (C) 2014 Elsevier Inc. All rights reserved.