Publications

Abstract

This chapter summarizes the extensive discussions that took place during the Forum as well as the subsequent months thereafter. It assesses current understanding of the neuronal mechanisms that underlie syntactic structure and processing.... It is posited that to understand the neurobiology of syntax, it might be worthwhile to shift the balance from comprehension to syntactic encoding in language production

Abstract

In this study, we investigate whether language and music share cognitive resources for structural processing. We report an experiment that used sung materials and manipulated linguistic complexity (subject-extracted relative clauses, object-extracted relative clauses) and musical complexity (in-key critical note, out-of-key critical note, auditory anomaly on the critical note involving a loudness increase). The auditory-anomaly manipulation was included in order to test whether the difference between in-key and out-of-key conditions might be due to any salient, unexpected acoustic event. The critical dependent measure involved comprehension accuracies to questions about the propositional content of the sentences asked at the end of each trial. The results revealed an interaction between linguistic and musical complexity such that the difference between the subject- and object-extracted relative clause conditions was larger in the out-of-key condition than in the in-key and auditory-anomaly conditions. These results provide evidence for an overlap in structural processing between language and music.

Abstract

The aim of the present work is to identify the evolutionary origins of the ability to speak and understand a natural language. I will adopt Botha’s “Windows Approach” (Language and Communication, 2006, 26, pp. 129–143) in order to justify the following two assumptions, which concern the evolutionary continuity between human language and animals’ communication systems: (a) despite the uniqueness of human language in sharing and conveying utterances with an open-ended structure, some isolated components of our linguistic competence are shared with non- human primates, grounding a line of evolutionary continuity; (b) the very first “linguistic” utterances were holistic, that is, whole bunches of sounds able to convey information despite their lack of modern syntax. I will address such suppositions through the comparative analysis of three constitutive features of human language: syntax, the semantic value of utterances, and the ability to attribute mental states to conspecifics, i.e. the theory of mind.

Abstract

Family and twin studies of developmental dyslexia have consistently shown that there is a significant heritable component for this disorder. However, any genetic basis for the trait is likely to be complex, involving reduced penetrance, phenocopy, heterogeneity and oligogenic inheritance. This complexity results in reduced power for traditional parametric linkage analysis, where specification of the correct genetic model is important. One strategy is to focus on large multigenerational pedigrees with severe phenotypes and/or apparent simple Mendelian inheritance, as has been successfully demonstrated for speech and language impairment. This approach is limited by the scarcity of such families. An alternative which has recently become feasible due to the development of high-throughput genotyping techniques is the analysis of large numbers of sib-pairs using allele-sharing methodology. This paper outlines our strategy for conducting a systematic genome-wide search for genes involved in dyslexia in a large number of affected sib-pair familites from the UK. We use a series of psychometric tests to obtain different quantitative measures of reading deficit, which should correlate with different components of the dyslexia phenotype, such as phonological awareness and orthographic coding ability. This enable us to use QTL (quantitative trait locus) mapping as a powerful tool for localising genes which may contribute to reading and spelling disability.

Abstract

Attention deficit/hyperactivity disorder (ADHD) is a common heritable disorder with a childhood onset. Molecular genetic studies of ADHD have previously focused on examining the roles of specific candidate genes, primarily those involved in dopaminergic pathways. We have performed the first systematic genomewide linkage scan for loci influencing ADHD in 126 affected sib pairs, using a ∼10-cM grid of microsatellite markers. Allele-sharing linkage methods enabled us to exclude any loci with a λs of ⩾3 from 96% of the genome and those with a λs of ⩾2.5 from 91%, indicating that there is unlikely to be a major gene involved in ADHD susceptibility in our sample. Under a strict diagnostic scheme we could exclude all screened regions of the X chromosome for a locus-specific λs of ⩾2 in brother-brother pairs, demonstrating that the excess of affected males with ADHD is probably not attributable to a major X-linked effect. Qualitative trait maximum LOD score analyses pointed to a number of chromosomal sites that may contain genetic risk factors of moderate effect. None exceeded genomewide significance thresholds, but LOD scores were >1.5 for regions on 5p12, 10q26, 12q23, and 16p13. Quantitative-trait analysis of ADHD symptom counts implicated a region on 12p13 (maximum LOD 2.6) that also yielded a LOD >1 when qualitative methods were used. A survey of regions containing 36 genes that have been proposed as candidates for ADHD indicated that 29 of these genes, including DRD4 and DAT1, could be excluded for a λs of 2. Only three of the candidates—DRD5, 5HTT, and CALCYON—coincided with sites of positive linkage identified by our screen. Two of the regions highlighted in the present study, 2q24 and 16p13, coincided with the top linkage peaks reported by a recent genome-scan study of autistic sib pairs.

Abstract

Recent application of nonparametric-linkage analysis to reading disability has implicated a putative quantitative-trait locus (QTL) on the short arm of chromosome 6. In the present study, we use QTL methods to evaluate linkage to the 6p25-21.3 region in a sample of 181 sib pairs from 82 nuclear families that were selected on the basis of a dyslexic proband. We have assessed linkage directly for several quantitative measures that should correlate with different components of the phenotype, rather than using a single composite measure or employing categorical definitions of subtypes. Our measures include the traditional IQ/reading discrepancy score, as well as tests of word recognition, irregular-word reading, and nonword reading. Pointwise analysis by means of sib-pair trait differences suggests the presence, in 6p21.3, of a QTL influencing multiple components of dyslexia, in particular the reading of irregular words (P=.0016) and nonwords (P=.0024). A complementary statistical approach involving estimation of variance components supports these findings (irregular words, P=.007; nonwords, P=.0004). Multipoint analyses place the QTL within the D6S422-D6S291 interval, with a peak around markers D6S276 and D6S105 consistently identified by approaches based on trait differences (irregular words, P=.00035; nonwords, P=.0035) and variance components (irregular words, P=.007; nonwords, P=.0038). Our findings indicate that the QTL affects both phonological and orthographic skills and is not specific to phoneme awareness, as has been previously suggested. Further studies will be necessary to obtain a more precise localization of this QTL, which may lead to the isolation of one of the genes involved in developmental dyslexia.

Abstract

Developmental dyslexia, a specific impairment of reading ability despite adequate intelligence and educational opportunity, is one of the most frequent childhood disorders. Since the first documented cases at the beginning of the last century, it has become increasingly apparent that the reading problems of people with dyslexia form part of a heritable neurobiological syndrome. As for most cognitive and behavioural traits, phenotypic definition is fraught with difficulties and the genetic basis is complex, making the isolation of genetic risk factors a formidable challenge. Against such a background, it is notable that several recent studies have reported the localization of genes that influence dyslexia and other language-related traits. These investigations exploit novel research approaches that are relevant to many areas of human neurogenetics.

Abstract

Rare mutations of the FOXP2 transcription factor gene cause a monogenic syndrome characterized by impaired speech development and linguistic deficits. Recent genomic investigations indicate that its downstream neural targets make broader impacts on common language impairments, bridging clinically distinct disorders. Moreover, the striking conservation of both FoxP2 sequence and neural expression in different vertebrates facilitates the use of animal models to study ancestral pathways that have been recruited towards human speech and language. Intriguingly, reduced FoxP2 dosage yields abnormal synaptic plasticity and impaired motor-skill learning in mice, and disrupts vocal learning in songbirds. Converging data indicate that Foxp2 is important for modulating the plasticity of relevant neural circuits. This body of research represents the first functional genetic forays into neural mechanisms contributing to human spoken language.

Abstract

Theories addressing the biological basis of language must be built on
an appreciation of the ways that molecular and neurobiological substrates
can contribute to aspects of human cognition. Here, we lay out
the principles by which a genome could potentially encode the necessary
information to produce a language-ready brain. We describe
what genes are; how they are regulated; and how they affect the formation,
function, and plasticity of neuronal circuits. At each step,
we give examples of molecules implicated in pathways that are important
for speech and language. Finally, we discuss technological advances
in genomics that are revealing considerable genotypic variation in
the human population, from rare mutations to common polymorphisms,
with the potential to relate this variation to natural variability
in speech and language skills. Moving forward, an interdisciplinary
approach to the language sciences, integrating genetics, neurobiology,
psychology, and linguistics, will be essential for a complete understanding
of our unique human capacities.

Abstract

Developmental dyslexia is defined as a specific and significant impairment in reading ability that cannot be explained by deficits in intelligence, learning opportunity, motivation or sensory acuity. It is one of the most frequently diagnosed disorders in childhood, representing a major educational and social problem. It is well established that dyslexia is a significantly heritable trait with a neurobiological basis. The etiological mechanisms remain elusive, however, despite being the focus of intensive multidisciplinary research. All attempts to map quantitative-trait loci (QTLs) influencing dyslexia susceptibility have targeted specific chromosomal regions, so that inferences regarding genetic etiology have been made on the basis of very limited information. Here we present the first two complete QTL-based genome-wide scans for this trait, in large samples of families from the United Kingdom and United States. Using single-point analysis, linkage to marker D18S53 was independently identified as being one of the most significant results of the genome in each scan (P< or =0.0004 for single word-reading ability in each family sample). Multipoint analysis gave increased evidence of 18p11.2 linkage for single-word reading, yielding top empirical P values of 0.00001 (UK) and 0.0004 (US). Measures related to phonological and orthographic processing also showed linkage at this locus. We replicated linkage to 18p11.2 in a third independent sample of families (from the UK), in which the strongest evidence came from a phoneme-awareness measure (most significant P value=0.00004). A combined analysis of all UK families confirmed that this newly discovered 18p QTL is probably a general risk factor for dyslexia, influencing several reading-related processes. This is the first report of QTL-based genome-wide scanning for a human cognitive trait.

Abstract

Between 2 and 5% of children who are otherwise unimpaired have significant difficulties in acquiring expressive and/or receptive language, despite adequate intelligence and opportunity. While twin studies indicate a significant role for genetic factors in developmental disorders of speech and language, the majority of families segregating such disorders show complex patterns of inheritance, and are thus not amenable for conventional linkage analysis. A rare exception is the KE family, a large three-generation pedigree in which approximately half of the members are affected with a severe speech and language disorder which appears to be transmitted as an autosomal dominant monogenic trait. This family has been widely publicised as suffering primarily from a defect in the use of grammatical suffixation rules, thus supposedly supporting the existence of genes specific to grammar. The phenotype, however, is broader in nature, with virtually every aspect of grammar and of language affected. In addition, affected members have a severe orofacial dyspraxia, and their speech is largely incomprehensible to the naive listener. We initiated a genome-wide search for linkage in the KE family and have identified a region on chromosome 7 which co-segregates with the speech and language disorder (maximum lod score = 6.62 at theta = 0.0), confirming autosomal dominant inheritance with full penetrance. Further analysis of microsatellites from within the region enabled us to fine map the locus responsible (designated SPCH1) to a 5.6-cM interval in 7q31, thus providing an important step towards its identification. Isolation of SPCH1 may offer the first insight into the molecular genetics of the developmental process that culminates in speech and language.

Abstract

This chapter highlights the research in isolating genetic factors underlying specific language impairment (SLI), or developmental dysphasia, which exploits newly developed genotyping technology, novel statistical methodology, and DNA sequence data generated by the Human Genome Project. The author begins with an overview of results from family, twin, and adoption studies supporting genetic involvement and then goes on to outline progress in a number of genetic mapping efforts that have been recently completed or are currently under way. It has been possible for genetic researchers to pinpoint the specific mutation responsible for some speech and language disorders, providing an example of how the availability of human genomic sequence data can greatly accelerate the pace of disease gene discovery. Finally, the author discusses future prospects on how molecular genetics may offer new insight into the etiology underlying speech and language disorders, leading to improvements in diagnosis and treatment.

Abstract

We present a neural-symbolic learning model of sentence production which displays strong semantic systematicity and recursive productivity. Using this model, we provide evidence for the data-driven learnability of complex yes/no- questions.

Abstract

The typological contrast between verb- and satellite-framed languages (Talmy, 1985) has set the basis for many empirical studies on L2 acquisition. The current analysis goes beyond this typology by looking in detail at the conceptualization of the path of motion in a motion event. We take as a starting point the cognitive salience of specific elements of motion events that are relevant when conceptualizing space. When expressing direction in French, specific spatial relations involving the entity in motion (its alignment and its distance toward a [potential] endpoint) are relevant, given a variety of path verbs in the lexicon expressing this information (e.g., se diriger vers, avancer to direct oneself toward,' to advance'). This is not the case in German (manner verbs in the lexicon mainly). In German, spatial information is packaged in adjuncts and particles and the path of motion is typically structured via features of the ground (entlanglaufen/fahren to walk/drive along') or endpoints (to walk/drive to/toward'). We investigate those fundamental differences in spatial conceptualization in French and German, as reflected in pre-articulatory patterns of attention allocation (measured with eye tracking) to moving entities and endpoints in motion scenes in an event description task. Our focus is on spatial conceptualization in an L2 (French L2 users of German), analyzing the extent to which these L2 users display target-like patterns or traces of L1 conceptualization transfer. Findings show that, in line with directional concepts expressed in verbs, L1 French speakers allocate more attention to entities in motion and endpoints, before utterance onset, than L1 German speakers do. The L2 German speakers pattern with L1 German speakers in the use of manner verbs, but they have not fully acquired the spatial concepts and means that structure the path of motion in the L2. This is reflected in pre-articulatory attention allocation patterns, according to which the L2 speakers pattern with native speakers of their L1 (French). The findings show a continued deep entrenchment of L1-based processing patterns and spatial frames of reference when speakers prepare for speech in an L2

Abstract

Studies on gaze allocation during sentence production have recently begun to implement cross-linguistic analyses in the investigation of visual and linguistic processing. The underlying assumption is that the aspects of a scene that attract attention prior to articulation are, in part, linked to the specifi c linguistic system and means used for expression. The present study concerns naturalistic, dynamic scenes (video clips) showing causative events (agent acting on an object) and exploits grammatical diff erences in the domain of verbal aspect, and the way in which the status of an event (a specifi c vs. habitual instance of an event) is encoded in English and German. Fixations in agent and action areas of interest were timelocked to utterance onset, and we focused on the pre-articulatory time span to shed light on sentence planning processes, involving message generation and scene conceptualization.

Abstract

We investigated whether brain potentials of grammatical aspect processing resemble semantic or morpho-syntactic processing, or whether they instead are characterized by an entirely distinct pattern in the same individuals. We studied aspect from the perspective of agreement between the temporal information in the context (temporal adverbials, e.g., Right now) and a morpho-syntactic marker of grammatical aspect (e.g., progressive is swimming). Participants read questions providing a temporal context that was progressive (What is Sophie doing in the pool right now?) or habitual (What does Sophie do in the pool every Monday?). Following a lead-in sentence context such as Right now, Sophie…, we measured event-related brain potentials (ERPs) time-locked to verb phrases in four different conditions, e.g., (a) is swimming (control); (b) ∗is cooking (semantic violation); (c) ∗are swimming (morpho-syntactic violation); or (d)?swims (aspect mismatch); …in the pool.” The collected ERPs show typical N400 and P600 effects for semantics and morpho-syntax, while aspect processing elicited an Early Negativity (250–350 ms). The aspect-related Negativity was short-lived and had a central scalp distribution with an anterior onset. This differentiates it not only from the semantic N400 effect, but also from the typical LAN (Left Anterior Negativity), that is frequently reported for various types of agreement processing. Moreover, aspect processing did not show a clear P600 modulation. We argue that the specific context for each item in this experiment provided a trigger for agreement checking with temporal information encoded on the verb, i.e., morphological aspect marking. The aspect-related Negativity obtained for aspect agreement mismatches reflects a violated expectation concerning verbal inflection (in the example above, the expected verb phrase was Sophie is X-ing rather than Sophie X-s in condition d). The absence of an additional P600 for aspect processing suggests that the mismatch did not require additional reintegration or processing costs. This is consistent with participants’ post hoc grammaticality judgements of the same sentences, which overall show a high acceptability of aspect mismatch sentences.

Abstract

Recent studies have identified neural correlates of language effects on perception in static domains of experience such as colour and objects. The generalization of such effects to dynamic domains like motion events remains elusive. Here, we focus on grammatical differences between languages relevant for the description of motion events and their impact on visual scene perception. Two groups of native speakers of German or English were presented with animated videos featuring a dot travelling along a trajectory towards a geometrical shape (endpoint). English is a language with grammatical aspect in which attention is drawn to trajectory and endpoint of motion events equally. German, in contrast, is a non-aspect language which highlights endpoints. We tested the comparative perceptual saliency of trajectory and endpoint of motion events by presenting motion event animations (primes) followed by a picture symbolising the event (target): In 75% of trials, the animation was followed by a mismatching picture (both trajectory and endpoint were different); in 10% of trials, only the trajectory depicted in the picture matched the prime; in 10% of trials, only the endpoint matched the prime; and in 5% of trials both trajectory and endpoint were matching, which was the condition requiring a response from the participant. In Experiment 1 we recorded event-related brain potentials elicited by the picture in native speakers of German and native speakers of English. German participants exhibited a larger P3 wave in the endpoint match than the trajectory match condition, whereas English speakers showed no P3 amplitude difference between conditions. In Experiment 2 participants performed a behavioural motion matching task using the same stimuli as those used in Experiment 1. German and English participants did not differ in response times showing that motion event verbalisation cannot readily account for the difference in P3 amplitude found in the first experiment. We argue that, even in a non-verbal context, the grammatical properties of the native language and associated sentence-level patterns of event encoding influence motion event perception, such that attention is automatically drawn towards aspects highlighted by the grammar.

Abstract

Los idiomas sudamericanas tienen una diversidad de sistemas numéricos, desde sistemas con solamente dos o tres términos en algunos idiomas amazónicos hasta sistemas con numerales extendiendo a miles. Una mirada al sistema del idioma cha’palaa de Ecuador demuestra rasgos de base-2, base-5, base-10 y base-20, ligados a diferentes etapas de cambio, desarrollo y contacto lingüístico. Conocer estas etapas nos permite proponer algunas correlaciones con lo que conocemos de la historia de contactos culturales en la región. The South American languages have diverse types of numeral systems, from systems of just two or three terms in some Amazonian languages to systems extending into the thousands. A look a the system of the Cha'palaa language of Ecuador demonstrates base-2, base-5, base-10 and base-20 features, linked to different stages of change, development and language contact. Learning about these stages permits up to propose some correlations between them and what we know about the history of cultural contact in the region.

Abstract

This article describes the interactional patterns and linguistic structures associated with otherinitiated repair, as observed in a corpus of video-recorded conversation in the Cha’palaa (a Barbacoan language spoken in north-western Ecuador). Special attention is given to the relation of repair formats to the morphosyntactic and intonational systems of the language. It examines the distinctive falling intonation observed with interjections and content question formats and the pattern of a held mid-high tone observed in polarity questions, as well as the function of Cha’palaa grammatical features such as the case marking system, the nominal classifiers and the verb classification system as formats for repair initiation. It considers a selection of examples from a video corpus to illustrate a broad range of sequence types of opened and restricted other-initiated repair, noting that Cha’palaa had the highest relative rate of open repair in the cross-linguistic sample. It also considers the extension of OIR to other practices such as news uptake and disagreement in the Cha’palaa corpus.

Abstract

The ethnographic literature has described the northwest Amazon as an area of shared culture across linguistic groups. This paper illustrates how a principle of semantic transparency across languages is a key means of establishing elements of a common regional culture through practices like the calquing of ethnonyms and toponyms so that they are semantically, but not phonologically, equivalent across languages. It places the northwest Amazon in a general discussion of cross-linguistic naming practices in South America and considers the extent to which a preference for semantic transparency can be linked to cases of widespread cultural “calquing”. It also addresses the principle of semantic transparency beyond specific referential phrases and into larger discourse structures. It concludes that an attention to semiotic practices in multilingual settings can provide new and more complex ways of thinking about the idea of shared culture

Abstract

In a 2x2 event-related FMRI study we find support for the idea that the inferior frontal cortex, centered on Broca’s region and its homologue, is involved in constructive unification operations during the structure-building process in parsing for comprehension. Tentatively, we provide evidence for a role of the dorsolateral prefrontal cortex centered on BA 9/46 in the control component of the language system. Finally, the left temporo-parietal cortex, in the vicinity of Wernicke’s region, supports the interaction between the syntax of gender agreement and sentence-level semantics.

Abstract

The nineteenth century witnessed some of the greatest neuroanatomists of all times. Amongst them is the largely forgotten Heinrich Sachs, a student of Carl Wernicke in Breslau.

Abstract

This is the first complete translation of Heinrich Sachs' outstanding white matter atlas dedicated to the occipital lobe. This work is accompanied by a prologue by Prof Carl Wernicke who for many years was Sachs' mentor in Breslau and enthusiastically supported his work.

Abstract

Human implicit learning can be investigated with implicit artificial grammar learning, a simple model for aspects of natural language acquisition. In this paper we investigate the remaining effect of modality transfer in syntactic classification of an acquired grammatical sequence structure after implicit grammar acquisition. Participants practiced either on acoustically presented syllable sequences or visually presented consonant letter sequences. During classification we independently manipulated the statistical frequency-based and rule-based characteristics of the classification stimuli. Participants performed reliably above chance on the within modality classification task although more so for those working on syllable sequence acquisition. These subjects were also the only group that kept a significant performance level in transfer classification. We speculate that this finding is of particular relevance in consideration of an ecological validity in the input signal in the use of artificial grammar learning and in language learning paradigms at large.

Abstract

Previous studies suggest that linguistic material can modulate visual perception, but it is unclear at which level of processing these interactions occur. Here we aim to dissociate between two competing models of language–perception interactions: a feed-forward and a feedback model. We capitalized on the fact that the models make different predictions on the role of feedback. We presented unmasked (aware) or masked (unaware) words implying motion (e.g. “rise,” “fall”), directly preceding an upward or downward visual motion stimulus. Crucially, masking leaves intact feed-forward information processing from low- to high-level regions, whereas it abolishes subsequent feedback. Under this condition, participants remained faster and more accurate when the direction implied by the motion word was congruent with the direction of the visual motion stimulus. This suggests that language–perception interactions are driven by the feed-forward convergence of linguistic and perceptual information at higher-level conceptual and decision stages.

Abstract

Previous studies have shown that language can modulate visual perception, by biasing and/
or enhancing perceptual performance. However, it is still debated where in the brain visual and
linguistic information are integrated, and whether the effects of language on perception are
automatic and persist even in the absence of awareness of the linguistic material. Here, we aimed
to explore the automaticity of language-perception interactions and the neural loci of these
interactions in an fMRI study. Participants engaged in a visual motion discrimination task (upward
or downward moving dots). Before each trial, a word prime was briefly presented that implied
upward or downward motion (e.g., “rise”, “fall”). These word primes strongly influenced behavior:
congruent motion words sped up reaction times and improved performance relative to incongruent
motion words. Neural congruency effects were only observed in the left middle temporal gyrus,
showing higher activity for congruent compared to incongruent conditions. This suggests that higherlevel
conceptual areas rather than sensory areas are the locus of language-perception interactions.
When motion words were rendered unaware by means of masking, they still affected visual motion
perception, suggesting that language-perception interactions may rely on automatic feed-forward
integration of perceptual and semantic material in language areas of the brain.

Abstract

Perception does not function as an isolated module but is tightly linked with other cognitive functions. Several studies have demonstrated an influence of language on motion perception, but it remains debated at which level of processing this modulation takes place. Some studies argue for an interaction in perceptual areas, but it is also possible that the interaction is mediated by "language areas" that integrate linguistic and visual information. Here, we investigated whether language-perception interactions were specific to the language-dominant left hemisphere by comparing the effects of language on visual material presented in the right (RVF) and left visual fields (LVF). Furthermore, we determined the neural locus of the interaction using fMRI. Participants performed a visual motion detection task. On each trial, the visual motion stimulus was presented in either the LVF or in the RVF, preceded by a centrally presented word (e.g., "rise"). The word could be congruent, incongruent, or neutral with regard to the direction of the visual motion stimulus that was presented subsequently. Participants were faster and more accurate when the direction implied by the motion word was congruent with the direction of the visual motion stimulus. Interestingly, the speed benefit was present only for motion stimuli that were presented in the RVF. We observed a neural counterpart of the behavioral facilitation effects in the left middle temporal gyrus, an area involved in semantic processing of verbal material. Together, our results suggest that semantic information about motion retrieved in language regions may automatically modulate perceptual decisions about motion.

Abstract

Genomewide quantitative-trait locus (QTL) linkage analysis was performed using a continuous measure of relative hand skill (PegQ) in a sample of 195 reading-disabled sibling pairs from the United Kingdom. This was the first genomewide screen for any measure related to handedness. The mean PegQ in the sample was equivalent to that of normative data, and PegQ was not correlated with tests of reading ability (correlations between −0.13 and 0.05). Relative hand skill could therefore be considered normal within the sample. A QTL on chromosome 2p11.2-12 yielded strong evidence for linkage to PegQ (empirical P=.00007), and another suggestive QTL on 17p11-q23 was also identified (empirical P=.002). The 2p11.2-12 locus was further analyzed in an independent sample of 143 reading-disabled sibling pairs, and this analysis yielded an empirical P=.13. Relative hand skill therefore is probably a complex multifactorial phenotype with a heterogeneous background, but nevertheless is amenable to QTL-based gene-mapping approaches.

Abstract

The molecular, developmental, and evolutionary bases of human brain asymmetry are almost completely unknown. Genetic linkage and association mapping have pin-pointed a gene called LRRTM1 (leucine-rich repeat transmembrane neuronal 1) that may contribute to variability in human handedness. Here I describe how LRRTM1's involvement in handedness was discovered, and also the latest knowledge of its functions in brain development and disease. The association of LRRTM1 with handedness was derived entirely from the paternally inherited gene, and follow-up analysis of gene expression confirmed that LRRTM1 is one of a small number of genes that are imprinted in the human genome, for which the maternally inherited copy is suppressed. The same variation at LRRTM1 that was associated paternally with mixed-/left-handedness was also over-transmitted paternally to schizophrenic patients in a large family study.
LRRTM1 is expressed in specific regions of the developing and adult forebrain by post-mitotic neurons, and the protein may be involved in axonal trafficking. Thus LRRTM1 has a probable role in neurodevelopment, and its association with handedness suggests that one of its functions may be in establishing or consolidating human brain asymmetry.
LRRTM1 is the first gene for which allelic variation has been associated with human handedness. The genetic data also suggest indirectly that the epigenetic regulation of this gene may yet prove more important than DNA sequence variation for influencing brain development and disease.
Intriguingly, the parent-of-origin activity of LRRTM1 suggests that men and women have had conflicting interests in relation to the outcome of lateralized brain development in their offspring.

Abstract

Lateralizations of brain structure and motor behavior have been observed in humans as early as the first trimester of gestation, and are likely to arise from asymmetrical genetic–developmental programs, as in other animals. Studies of gene expression levels in postmortem tissue samples, comparing the left and right sides of the human cerebral cortex, have generally not revealed striking transcriptional differences between the hemispheres. This is likely due to lateralization of gene expression being subtle and quantitative. However, a recent re-analysis and meta-analysis of gene expression data from the adult superior temporal and auditory cortex found lateralization of transcription of genes involved in synaptic transmission and neuronal electrophysiology. Meanwhile, human subcortical mid- and hindbrain structures have not been well studied in relation to lateralization of gene activity, despite being potentially important developmental origins of asymmetry. Genetic polymorphisms with small effects on adult brain and behavioral asymmetries are beginning to be identified through studies of large datasets, but the core genetic mechanisms of lateralized human brain development remain unknown. Identifying subtly lateralized genetic networks in the brain will lead to a new understanding of how neuronal circuits on the left and right are differently fine-tuned to preferentially support particular cognitive and behavioral functions.

Abstract

A locus on chromosome 2p12-16 has been implicated in dyslexia susceptibility by two independent linkage studies, including our own study of 119 nuclear twin-based families, each with at least one reading-disabled child. Nonetheless, no variant of any gene has been reported to show association with dyslexia, and no consistent clinical evidence exists to identify candidate genes with any strong a priori logic. We used 21 microsatellite markers spanning 2p12-16 to refine our 1-LOD unit linkage support interval to 12cM between D2S337 and D2S286. Then, in quantitative association analysis, two microsatellites yielded P values<0.05 across a range of reading-related measures (D2S2378 and D2S2114). The exon/intron borders of two positional candidate genes within the region were characterized, and the exons were screened for polymorphisms. The genes were Semaphorin4F (SEMA4F), which encodes a protein involved in axonal growth cone guidance, and OTX1, encoding a homeodomain transcription factor involved in forebrain development. Two non-synonymous single nucleotide polymorphisms were found in SEMA4F, each with a heterozygosity of 0.03. One intronic single nucleotide polymorphism between exons 12 and 13 of SEMA4F was tested for quantitative association, but no significant association was found. Only one single nucleotide polymorphism was found in OTX1, which was exonic but silent. Our data therefore suggest that linkage with reading disability at 2p12-16 is not caused by coding variants of SEMA4F or OTX1. Our study outlines the approach necessary for the identification of genetic variants causing dyslexia susceptibility in an epidemiological population of dyslexics.

Abstract

Dyslexia, a disorder of reading and spelling, is a heterogeneous neurological syndrome with a complex genetic and environmental aetiology. People with dyslexia differ in their individual profiles across a range of cognitive, physiological, and behavioural measures related to reading disability. Some or all of the subtypes of dyslexia might have partly or wholly distinct genetic causes. An understanding of the role of genetics in dyslexia could help to diagnose and treat susceptible children more effectively and rapidly than is currently possible and in ways that account for their individual disabilities. This knowledge will also give new insights into the neurobiology of reading and language cognition. Genetic linkage analysis has identified regions of the genome that might harbour inherited variants that cause reading disability. In particular, loci on chromosomes 6 and 18 have shown strong and replicable effects on reading abilities. These genomic regions contain tens or hundreds of candidate genes, and studies aimed at the identification of the specific causal genetic variants are underway.

Abstract

Francks et al. (2007) performed a recent study in which the first putative genetic effect on human handedness was identified (the imprinted locus LRRTM1 on human chromosome 2). In this issue of Laterality, Tim Crow and colleagues present a critique of that study. The present paper presents a personal response to that critique which argues that Francks et al. (2007) published a substantial body of evidence implicating LRRTM1 in handedness and schizophrenia. Progress will now be achieved by others trying to validate, refute, or extend those findings, rather than by further armchair discussion.

Abstract

This study aims to test a prediction of recent
theoretical frameworks in speech motor control: if speech production targets are specified in auditory
terms, people with better auditory acuity should have more precise speech targets.
To investigate this, we had participants perform speech perception and production tasks in a counterbalanced order. To assess speech perception acuity, we used an adaptive speech discrimination
task. To assess variability in speech production, participants performed a pseudo-word reading task; formant values were measured for each recording.
We predicted that speech production variability to correlate inversely with discrimination performance.
The results suggest that people do vary in their production and perceptual abilities, and that better discriminators have more distinctive vowel production targets, confirming our prediction. This
study highlights the importance of individual
differences in the study of speech motor control, and sheds light on speech production-perception interaction.

Abstract

Models of speech production explain event-related suppression of the auditory cortical
response as reflecting a comparison between auditory predictions and feedback. The present MEG
study was designed to test two predictions from this framework: 1) whether the reduced auditory
response varies as a function of the mismatch between prediction and feedback; 2) whether individual
variation in this response is predictive of speech-motor adaptation.
Participants alternated between online imitation and listening tasks. In the imitation task, participants
began each trial producing the same vowel (/e/) and subsequently listened to and imitated auditorilypresented
vowels varying in acoustic distance from /e/.
Results replicated suppression, with a smaller M100 during speaking than listening. Although we did
not find unequivocal support for the first prediction, participants with less M100 suppression were
better at the imitation task. These results are consistent with the enhancement of M100 serving as an
error signal to drive subsequent speech-motor adaptation.

Abstract

Theeffects of spatial filtering in functional magnetic resonance imaging were investigated by reevaluating the data of a previous study of episodic memory encoding at 2 × 2 × 4-mm3 resolution with use of a SPM99 analysis involving a Gaussian kernel of 8-mm full width at half maximum. In addition, a multisubject analysis of activated regions was performed by normalizing the functional images to an approximate Talairach brain atlas. In individual subjects, spatial filtering merged activations in anatomically separated brain regions. Moreover, small foci of activated pixels which originated from veins became blurred and hence indistinguishable from parenchymal responses. The multisubject analysis resulted in activation of the hippocampus proper, a finding which could not be confirmed by the activation maps obtained at high resolution. It is concluded that the validity of multisubject fMRI analyses can be considerably improved by first analyzing individual data sets at optimum resolution to assess the effects of spatial filtering and minimize the risk of signal contamination by macroscopically visible vessels.

Abstract

PTEN is a tumor suppressor associated with an inherited cancer syndrome and an important regulator of ongoing neural connectivity and plasticity. The present study examined molecular and phenotypic characteristics of individuals with germline heterozygous PTEN mutations and autism spectrum disorder (ASD) (PTEN-ASD), with the aim of identifying pathophysiologic markers that specifically associate with PTEN-ASD and that may serve as targets for future treatment trials. PTEN-ASD patients (n=17) were compared with idiopathic (non-PTEN) ASD patients with (macro-ASD, n=16) and without macrocephaly (normo-ASD, n=38) and healthy controls (n=14). Group differences were evaluated for PTEN pathway protein expression levels, global and regional structural brain volumes and cortical thickness measures, neurocognition and adaptive behavior. RNA expression patterns and brain characteristics of a murine model of Pten mislocalization were used to further evaluate abnormalities observed in human PTEN-ASD patients. PTEN-ASD had a high proportion of missense mutations and showed reduced PTEN protein levels. Compared with the other groups, prominent white-matter and cognitive abnormalities were specifically associated with PTEN-ASD patients, with strong reductions in processing speed and working memory. White-matter abnormalities mediated the relationship between PTEN protein reductions and reduced cognitive ability. The Ptenm3m4 murine model had differential expression of genes related to myelination and increased corpus callosum. Processing speed and working memory deficits and white-matter abnormalities may serve as useful features that signal clinicians that PTEN is etiologic and prompting referral to genetic professionals for gene testing, genetic counseling and cancer risk management; and could reveal treatment targets in trials of treatments for PTEN-ASD.

Abstract

The past two decades have seen an upsurge of interest in the collective behaviors of complex systems composed of many agents entrained to each other and to external events. In this paper, we extend the concept of entrainment to the dynamics of human collective attention. We conducted a detailed investigation of the unfolding of human entrainment—as expressed by the content and patterns of hundreds of thousands of messages on Twitter—during the 2012 US presidential debates. By time-locking these data sources, we quantify the impact of the unfolding debate on human attention at three time scales. We show that collective social behavior covaries second-by-second to the interactional dynamics of the debates: A candidate speaking induces rapid increases in mentions of his name on social media and decreases in mentions of the other candidate. Moreover, interruptions by an interlocutor increase the attention received. We also highlight a distinct time scale for the impact of salient content during the debates: Across well-known remarks in each debate, mentions in social media start within 5–10 seconds after it occurs; peak at approximately one minute; and slowly decay in a consistent fashion across well-known events during the debates. Finally, we show that public attention after an initial burst slowly decays through the course of the debates. Thus we demonstrate that large-scale human entrainment may hold across a number of distinct scales, in an exquisitely time-locked fashion. The methods and results pave the way for careful study of the dynamics and mechanisms of large-scale human entrainment.

Abstract

Photosensitivity is a heritable abnormal cortical response to flickering light, manifesting as particular electroencephalographic changes, with or without seizures. Photosensitivity is prominent in a very rare epileptic encephalopathy due to de novo CHD2 mutations, but is also seen in epileptic encephalopathies due to other gene mutations. We determined whether CHD2 variation underlies photosensitivity in common epilepsies, specific photosensitive epilepsies and individuals with photosensitivity without seizures. We studied 580 individuals with epilepsy and either photosensitive seizures or abnormal photoparoxysmal response on electroencephalography, or both, and 55 individuals with photoparoxysmal response but no seizures. We compared CHD2 sequence data to publicly available data from 34 427 individuals, not enriched for epilepsy. We investigated the role of unique variants seen only once in the entire data set. We sought CHD2 variants in 238 exomes from familial genetic generalized epilepsies, and in other public exome data sets. We identified 11 unique variants in the 580 individuals with photosensitive epilepsies and 128 unique variants in the 34 427 controls: unique CHD2 variation is over-represented in cases overall (P = 2·17 × 10−5). Among epilepsy syndromes, there was over-representation of unique CHD2 variants (3/36 cases) in the archetypal photosensitive epilepsy syndrome, eyelid myoclonia with absences (P = 3·50 × 10−4). CHD2 variation was not over-represented in photoparoxysmal response without seizures. Zebrafish larvae with chd2 knockdown were tested for photosensitivity. Chd2 knockdown markedly enhanced mild innate zebrafish larval photosensitivity. CHD2 mutation is the first identified cause of the archetypal generalized photosensitive epilepsy syndrome, eyelid myoclonia with absences. Unique CHD2 variants are also associated with photosensitivity in common epilepsies. CHD2 does not encode an ion channel, opening new avenues for research into human cortical excitability.

Abstract

In this paper we present the first results of the application of computational methods, inspired by the ideas in McMahon & McMahon (2005), to a dataset collected from languages of every branch of the Tupian family (including all living non-Tupí-Guaraní languages) in order to produce a classification of the family based on lexical distance. We used both a Swadesh list (with historically stabler terms) and a list of animal and plant names for results comparison. In addition, we also selected more (HiHi) and less (LoLo) stable terms from the Swadesh list to form sublists for indepedent treatment. We compared the resulting NeighborNet networks and neighbor-joining cladograms and drew conclusions about their significance for the current understanding of the classification of Tupian languages. One important result is the lack of support for the currently discussed idea of an Eastern-Western division within Tupí

Abstract

This study addresses how verbal self-monitoring and the Error-Related Negativity (ERN) are affected by time pressure
when a task is performed in a second language as opposed to performance in the native language. German–Dutch
bilinguals were required to perform a phoneme-monitoring task in Dutch with and without a time pressure manipulation.
We obtained an ERN following verbal errors that showed an atypical increase in amplitude under time
pressure. This finding is taken to suggest that under time pressure participants had more interference from their native
language, which in turn led to a greater response conflict and thus enhancement of the amplitude of the ERN. This
result demonstrates once more that the ERN is sensitive to psycholinguistic manipulations and suggests that the
functioning of the verbal self-monitoring systemduring speaking is comparable to other performance monitoring, such
as action monitoring.

Abstract

This paper presents a new architecture for optical burst switched networks where the control plane of the network functions in a cooperative manner. Each node interprets the data conveyed by the control packet and forwards it to the next nodes, making the control plane of the network distribute the relevant information to all the nodes in the network. A cooperation transmission tree is used, thus allowing all the nodes to store the information related to the traffic management in the network, and enabling better network resource planning at each node. A model of this network architecture is proposed, and its performance is evaluated.

Abstract

Phonagnosia, the inability to recognize familiar voices, has been studied in brain-damaged patients but no cases due to developmental problems have been reported. Here we describe the case of KH, a 60-year-old active professional woman who reports that she has always experienced severe voice recognition difficulties. Her hearing abilities are normal, and an MRI scan showed no evidence of brain damage in regions associated with voice or auditory perception. To better understand her condition and to assess models of voice and high-level auditory processing, we tested KH on behavioural tasks measuring voice recognition, recognition of vocal emotions, face recognition, speech perception, and processing of environmental sounds and music. KH was impaired on tasks requiring the recognition of famous voices and the learning and recognition of new voices. In contrast, she performed well on nearly all other tasks. Her case is the first report of developmental phonagnosia, and the results suggest that the recognition of a speaker’s vocal identity depends on separable mechanisms from those used to recognize other information from the voice or non-vocal auditory stimuli.

Abstract

Mutations of the forkhead transcription factor FOXP2 gene have been implicated in inherited speech-and-language disorders, and specific Foxp2 expression patterns in neuronal populations and neuronal phenotypes arising from Foxp2 disruption have been described. However, molecular functions of FOXP2 are not completely understood. Here we report a requirement for FOXP2 in growth arrest of the osteosarcoma cell line 143B. We observed endogenous expression of this transcription factor both transiently in normally developing murine osteoblasts and constitutively in human SAOS-2 osteosarcoma cells blocked in early osteoblast development. Critically, we demonstrate that in 143B osteosarcoma cells with minimal endogenous expression, FOXP2 induced by growth arrest is required for up-regulation of p21WAF1/CIP1. Upon growth factor withdrawal, FOXP2 induction occurs rapidly and precedes p21WAF1/CIP1 activation. Additionally, FOXP2 expression could be induced by MAPK pathway inhibition in growth-arrested 143B cells, but not in traditional cell line models of osteoblast differentiation (MG-63, C2C12, MC3T3-E1). Our data are consistent with a model in which transient upregulation of Foxp2 in pre-osteoblast mesenchymal cells regulates a p21-dependent growth arrest checkpoint, which may have implications for normal mesenchymal and osteosarcoma biology

Abstract

In the context of large and ever growing archives, generating annotation suggestions automatically from textual resources related to the documents to be archived is an interesting option in theory. It could save a lot of work in the time consuming and expensive task of manual annotation and it could help cataloguers attain a higher inter-annotator agreement. However, some questions arise in practice: what is the quality of the automatically produced annotations? How do they compare with manual annotations and with the requirements for annotation that were defined in the archive? If different from the manual annotations, are the automatic annotations wrong? In the CHOICE project, partially hosted at the Netherlands Institute for Sound and Vision, the Dutch public archive for audiovisual broadcasts, we automatically generate annotation suggestions for cataloguers. In this paper, we define three types of evaluation of these annotation suggestions: (1) a classic and strict evaluation measure expressing the overlap between automatically generated keywords and the manual annotations, (2) a loosened evaluation measure for which semantically very similar annotations are also considered as relevant matches, and (3) an in-use evaluation of the usefulness of manual versus automatic annotations in the context of serendipitous browsing. During serendipitous browsing, the annotations (manual or automatic) are used to retrieve and visualize semantically related documents.

Abstract

Importance 
Neuroticism is a pervasive risk factor for psychiatric conditions. It genetically overlaps with major depressive disorder (MDD) and is therefore an important phenotype for psychiatric genetics. The Genetics of Personality Consortium has created a resource for genome-wide association analyses of personality traits in more than 63 000 participants (including MDD cases).Objectives
To identify genetic variants associated with neuroticism by performing a meta-analysis of genome-wide association results based on 1000 Genomes imputation; to evaluate whether common genetic variants as assessed by single-nucleotide polymorphisms (SNPs) explain variation in neuroticism by estimating SNP-based heritability; and to examine whether SNPs that predict neuroticism also predict MDD.Design, Setting, and Participants
Genome-wide association meta-analysis of 30 cohorts with genome-wide genotype, personality, and MDD data from the Genetics of Personality Consortium. The study included 63 661 participants from 29 discovery cohorts and 9786 participants from a replication cohort. Participants came from Europe, the United States, or Australia. Analyses were conducted between 2012 and 2014.Main Outcomes and Measures
Neuroticism scores harmonized across all 29 discovery cohorts by item response theory analysis, and clinical MDD case-control status in 2 of the cohorts.Results
A genome-wide significant SNP was found on 3p14 in MAGI1 (rs35855737; P = 9.26 × 10−9 in the discovery meta-analysis). This association was not replicated (P = .32), but the SNP was still genome-wide significant in the meta-analysis of all 30 cohorts (P = 2.38 × 10−8). Common genetic variants explain 15% of the variance in neuroticism. Polygenic scores based on the meta-analysis of neuroticism in 27 cohorts significantly predicted neuroticism (1.09 × 10−12 <} P {<} .05) and MDD (4.02 × 10−9 {<} P {< .05) in the 2 other cohorts.Conclusions and Relevance
This study identifies a novel locus for neuroticism. The variant is located in a known gene that has been associated with bipolar disorder and schizophrenia in previous studies. In addition, the study shows that neuroticism is influenced by many genetic variants of small effect that are either common or tagged by common variants. These genetic variants also influence MDD. Future studies should confirm the role of the MAGI1 locus for neuroticism and further investigate the association of MAGI1 and the polygenic association to a range of other psychiatric disorders that are phenotypically correlated with neuroticism.

Abstract

The acquisition of linguistic structures that require perspective-taking at the level of message generation is challenging. We investigate use of progressive aspect in L2 event encoding, using a sentence priming paradigm. We focus on Dutch, in which use of progressive aspect is optional. The progressive consists of a prepositional phrase (‘aan het,’ at-the), plus a verbal infinitive. We ask, to what extent L2 speakers, in comparison to native speakers, show priming effects in relation to form (prepositional phrase) or conceptual (progressive aspect) prime sentences. In native Dutch speakers we find a priming effect for the ‘progressive prime,’ compared to a ‘neutral prime’ (aspectually neutral event description). In L2 speakers this effect was absent. For the form prime, no priming effects were obtained in native speakers, rather, we find evidence for a partial blocking effect in L2 speakers. Results suggest that the strength of the link between concept and form of progressive aspect differs in native and L2 speakers. Specific factors contributed to the L2 findings, e.g., level of L2 proficiency and degree of L2 exposure. We conclude that (1) the conceptual basis of grammatical aspect can be primed in native speakers, and (2) in L2 speakers, access to conceptual information is less automatized.

Abstract

Gricean pragmatics has often been criticised for being implausible from a psychological point of view. This line of criticism is never backed up by empirical evidence, but more importantly, it ignores the fact that Grice never meant to advance a processing theory, in the first place. Taking our lead from Marr (1982), we distinguish between two levels of explanation: at the W-level, we are concerned with what agents do and why; at the H-level, we ask how agents do whatever it is they do. Whereas pragmatics is pitched at the W-level, processing theories are at the H-level. This is not to say that pragmatics has no implications for psychology at all, but it is to say that its implications are less direct than is often supposed.

Abstract

A key function of attention is to select an appropriate subset of available information by facilitation of attended processes and/or inhibition of irrelevant processing. Functional imaging studies, using positron emission tomography, have during different experimental tasks revealed decreased neuronal activity in areas that process input from unattended sensory modalities. It has been hypothesized that these decreases reflect a selective inhibitory modulation of nonrelevant cortical processing. In this study we addressed this question using a continuous arithmetical task with and without concomitant disturbing auditory input (task-irrelevant speech). During the arithmetical task, irrelevant speech did not affect task-performance but yielded decreased activity in the auditory and midcingulate cortices and increased activity in the left posterior parietal cortex. This pattern of modulation is consistent with a top down inhibitory modulation of a nonattended input to the auditory cortex and a coexisting, attention-based facilitation of taskrelevant processing in higher order cortices. These findings suggest that task-related decreases in cortical activity may be of functional importance in the understanding of both attentional mechanisms and taskrelated information processing.

Abstract

In cochlear implants (CIs), acoustic speech cues, especially for pitch, are delivered in a degraded form. This study's aim is to assess whether due to degraded pitch cues, normal-hearing listeners and CI users employ different perceptual strategies to recognize vocal emotions, and, if so, how these differ. Voice actors were recorded pronouncing a nonce word in four different emotions: anger, sadness, joy, and relief. These recordings' pitch cues were phonetically analyzed. The recordings were used to test 20 normal-hearing listeners' and 20 CI users' emotion recognition. In congruence with previous studies, high-arousal emotions had a higher mean pitch, wider pitch range, and more dominant pitches than low-arousal emotions. Regarding pitch, speakers did not differentiate emotions based on valence but on arousal. Normal-hearing listeners outperformed CI users in emotion recognition, even when presented with CI simulated stimuli. However, only normal-hearing listeners recognized one particular actor's emotions worse than the other actors'. The groups behaved differently when presented with similar input, showing that they had to employ differing strategies. Considering the respective speaker's deviating pronunciation, it appears that for normal-hearing listeners, mean pitch is a more salient cue than pitch range, whereas CI users are biased toward pitch range cues

Abstract

Advances in molecular technologies make it possible to pinpoint genomic factors associated with complex human traits. For cognition and behaviour, identification of underlying genes provides new entry points for deciphering the key neurobiological pathways. In the past decade, the search for genetic correlates of musicality has gained traction. Reports have documented familial clustering for different extremes of ability, including amusia and absolute pitch (AP), with twin studies demonstrating high heritability for some music-related skills, such as pitch perception. Certain chromosomal regions have been linked to AP and musical aptitude, while individual candidate genes have been investigated in relation to aptitude and creativity. Most recently, researchers in this field started performing genome-wide association scans. Thus far, studies have been hampered by relatively small sample sizes and limitations in defining components of musicality, including an emphasis on skills that can only be assessed in trained musicians. With opportunities to administer standardized aptitude tests online, systematic large-scale assessment of musical abilities is now feasible, an important step towards high-powered genome-wide screens. Here, we offer a synthesis of existing literatures and outline concrete suggestions for the development of comprehensive operational tools for the analysis of musical phenotypes.

Abstract

The ability to recognize speech acts (verbal actions) in conversation is critical for everyday interaction. However, utterances are often underspecified for the speech act they perform, requiring listeners to rely on the context to recognize the action. The goal of this study was to investigate the time-course of auditory speech act recognition in action-underspecified utterances and explore how sequential context (the prior action) impacts this process. We hypothesized that speech acts are recognized early in the utterance to allow for quick transitions between turns in conversation. Event-related potentials (ERPs) were recorded while participants listened to spoken dialogues and performed an action categorization task. The dialogues contained target utterances that each of which could deliver three distinct speech acts depending on the prior turn. The targets were identical across conditions, but differed in the type of speech act performed and how it fit into the larger action sequence. The ERP results show an early effect of action type, reflected by frontal positivities as early as 200 ms after target utterance onset. This indicates that speech act recognition begins early in the turn when the utterance has only been partially processed. Providing further support for early speech act recognition, actions in highly constraining contexts did not elicit an ERP effect to the utterance-final word. We take this to show that listeners can recognize the action before the final word through predictions at the speech act level. However, additional processing based on the complete utterance is required in more complex actions, as reflected by a posterior negativity at the final word when the speech act is in a less constraining context and a new action sequence is initiated. These findings demonstrate that sentence comprehension in conversational contexts crucially involves recognition of verbal action which begins as soon as it can.

Abstract

The ability to repair problems with hearing or understanding in conversation is critical for successful communication. This article describes the linguistic practices of other-initiated repair (OIR) in Icelandic through quantitative and qualitative analysis of a corpus of video-recorded conversations. The study draws on the conceptual distinctions developed in the comparative project on repair described in the introduction to this issue. The main aim is to give an overview of the formats for OIR in Icelandic and the type of repair practices engendered by them. The use of repair initiations in social actions not aimed at solving comprehension problems is also briefly discussed. In particular, the interjection ha has a rich usage extending beyond open other-initiation of repair. By describing the linguistic machinery for other-initiated repair in Icelandic, this study contributes to the typology of conversational structure and to the still nascent field of Icelandic social interaction studies.

Abstract

OBJECTIVES: Combining the analysis of family-based samples with unrelated individuals can enhance the power of genetic association studies. Various combined analysis techniques have been recently developed; as yet, there have been no comparisons of their power, or robustness to confounding factors. We investigated empirically the power of up to six combined methods using simulated samples of trios and unrelated cases/controls (TDTCC), trios and unrelated controls (TDTC), and affected sibpairs with parents and unrelated cases/controls (ASPFCC). METHODS: We simulated multiplicative, dominant and recessive models with varying risk parameters in single samples. Additionally, we studied false-positive rates and investigated, if possible, the coverage of the true genetic effect (TDTCC). RESULTS/CONCLUSIONS: Under the TDTCC design, we identified four approaches with equivalent power and false-positive rates. Combined statistics were more powerful than single-sample statistics or a pooled chi(2)-statistic when risk parameters were similar in single samples. Adding parental information to the CC part of the joint likelihood increased the power of generalised logistic regression under the TDTC but not the TDTCC scenario. Formal testing of differences between risk parameters in subsamples was the most sensitive approach to avoid confounding in combined analysis. Non-parametric analysis based on Monte-Carlo testing showed the highest power for ASPFCC samples.

Abstract

The verb has traditionally been characterized as the central element in a sentence. Nevertheless, the exact role of the verb during the actual ongoing comprehension of a sentence as it unfolds in time remains largely unknown. This paper reports the results of two Cross-Modal Lexical Priming (CMLP) experiments detailing the pattern of verb priming during on-line processing of Dutch sentences. Results are contrasted with data from a third CMLP experiment on priming of nouns in similar sentences. It is demonstrated that the meaning of a matrix verb remains active throughout the entire matrix clause, while this is not the case for the meaning of a subject head noun. Activation of the meaning of the verb only dissipates upon encountering a clear signal as to the start of a new clause.

Abstract

Deaf children whose hearing losses prevent them from accessing spoken language and whose hearing parents have not exposed them to sign language develop gesture systems, called homesigns, which have many of the properties of natural language—the so-called resilient properties of language. We explored the resilience of structure built around the predicate—in particular, how manner and path are mapped onto the verb—in homesign systems developed by deaf children in Turkey and the United States. We also asked whether the Turkish homesigners exhibit sentence-level structures previously identified as resilient in American and Chinese homesigners. We found that the Turkish and American deaf children used not only the same production probability and ordering patterns to indicate who does what to whom, but also used the same segmentation and conflation patterns to package manner and path. The gestures that the hearing parents produced did not, for the most part, display the patterns found in the children's gestures. Although cospeech gesture may provide the building blocks for homesign, it does not provide the blueprint for these resilient properties of language.

Abstract

Vocal individuality provides a method of personalization for multiple avian species. However, expression of individual vocal features depends on necessity of recognition. Here we focused on chick vocalizations of demoiselle, Siberian and red-crowned cranes that differ by their body size, developmental rates and some ecological traits. Cranes are territorial during summer, but gather in
large flocks during autumn and winter. Nevertheless, parents keep feeding their chicks, even on winter grounds, despite the potential of confusing their own and alien
chicks. Here we aimed to compare expression of individuality and sex in calls of three crane species between solitary and gregarious periods of a chick’s life, and between species. We found significant individual patterns of
acoustic variables in the calls of all three species both before and after fledging. However, only red-crowned crane chicks increased expression of individuality significantly after the fledging. Also, we found that chicks of all three species significantly increased occurrence of nonlinear phenomena, i.e., irregular oscillations of soundproducing membranes (biphonations, sidebands, and deterministic chaos), in their calls after fledging. Non-linear phenomena can be a way of increasing the potential for
individual recognition as well as avoiding habituation of parents to their chicks’ calls. The older chicks are, the less
their parents feed them, and chicks benefit from keeping the permanent attention.

Abstract

Learning to recognize the contrasts of a language-specific phonemic repertoire can be viewed as forming categories in a multidimensional psychophysical space. Research on the learning of distributionally defined visual categories has shown that categories defined over I dimension are easy to learn and that learning multidimensional categories is more difficult but tractable under specific task conditions. In 2 experiments, adult participants learned either a unidimensional ora multidimensional category distinction with or without supervision (feedback) during learning. The unidimensional distinctions were readily learned and supervision proved beneficial, especially in maintaining category learning beyond the learning phase. Learning the multidimensional category distinction proved to be much more difficult and supervision was not nearly as beneficial as with unidimensionally defined categories. Maintaining a learned multidimensional category distinction was only possible when the distributional information (hat identified the categories remained present throughout the testing phase. We conclude that listeners are sensitive to both trial-by-trial feedback and the distributional information in the stimuli. Even given limited exposure, listeners learned to use 2 relevant dimensions. albeit with considerable difficulty.

Abstract

The human capacity to acquire sophisticated language is unmatched in the animal kingdom. Despite the discontinuity in communicative abilities between humans and other primates, language is built on ancient genetic foundations, which are being illuminated by comparative genomics. The genetic architecture of the language faculty is also being uncovered by research into neurodevelopmental disorders that disrupt the normally effortless process of language acquisition. In this article, we discuss the strategies that researchers are using to reveal genetic factors contributing to communicative abilities, and review progress in identifying the relevant genes and genetic variants. The first gene directly implicated in a speech and language disorder was FOXP2. Using this gene as a case study, we illustrate how evidence from genetics, molecular cell biology, animal models and human neuroimaging has converged to build a picture of the role of FOXP2 in neurodevelopment, providing a framework for future endeavors to bridge the gaps between genes, brains and behavior

Abstract

Prolectin, a previously undescribed glycan-binding receptor, has been identified by re-screening of the human genome for genes encoding proteins containing potential C-type carbohydrate-recognition domains. Glycan array analysis revealed that the carbohydrate-recognition domain in the extracellular domain of the receptor binds glycans with terminal α-linked mannose or fucose residues. Prolectin expressed in fibroblasts is found at the cell surface, but unlike many glycan-binding receptors it does not mediate endocytosis of a neoglycoprotein ligand. However, compared with other known glycan-binding receptors, the receptor contains an unusually large intracellular domain that consists of multiple sequence motifs, including phosphorylated tyrosine residues, that allow it to interact with signaling molecules such as Grb2. Immunohistochemistry has been used to demonstrate that prolectin is expressed on a specialized population of proliferating B cells in germinal centers. Thus, this novel receptor has the potential to function in carbohydrate-mediated communication between cells in the germinal center.

Abstract

Language is a defining characteristic of the human species, but its foundations remain mysterious. Heritable disorders offer a gateway into biological underpinnings, as illustrated by the discovery that FOXP2 disruptions cause a rare form of speech and language impairment. The genetic architecture underlying language-related disorders is complex, and although some progress has been made, it has proved challenging to pinpoint additional relevant genes with confidence. Next-generation sequencing and genome-wide association studies are revolutionizing understanding of the genetic bases of other neurodevelopmental disorders, like autism and schizophrenia, and providing fundamental insights into the molecular networks crucial for typical brain development. We discuss how a similar genomic perspective, brought to the investigation of language-related phenotypes, promises to yield equally informative discoveries. Moreover, we outline how follow-up studies of genetic findings using cellular systems and animal models can help to elucidate the biological mechanisms involved in the development of brain circuits supporting language.

Abstract

The current study investigated whether point-accompanying characteristics, like vocalizations and hand shape, differentiate infants' underlying motives of prelinguistic pointing. We elicited imperative (requestive) and declarative (expressive and informative) pointing acts in experimentally controlled situations, and analyzed accompanying characteristics. Experiment 1 revealed that prosodic characteristics of point-accompanying vocalizations distinguished requestive from both expressive and informative pointing acts, with little differences between the latter two. In addition, requestive points were more often realized with the whole hand than the index finger, while this was the opposite for expressive and informative acts. Experiment 2 replicated Experiment 1, revealing distinct prosodic characteristics for requestive pointing also when the referent was distal and when it had an index-finger shape. Findings reveal that beyond the social context, point-accompanying vocalizations give clues to infants' underlying intentions when pointing.

Abstract

The genetic determinants of cerebral asymmetries are unknown. Sex differences in asymmetry of the planum temporale, that overlaps Wernicke’s classical language area, have been inconsistently reported. Meta-analysis of previous studies has suggested that publication bias established this sex difference in the literature. Using probabilistic definitions of cortical regions we screened over the cerebral cortex for sexual dimorphisms of asymmetry in 2337 healthy subjects, and found the planum temporale to show the strongest sex-linked asymmetry of all regions, which was supported by two further datasets, and also by analysis with the Freesurfer package that performs automated parcellation of cerebral cortical regions. We performed a genome-wide association scan meta-analysis of planum temporale asymmetry in a pooled sample of 3095 subjects, followed by a candidate-driven approach which measured a significant enrichment of association in genes of the ´steroid hormone receptor activity´ and 'steroid metabolic process' pathways. Variants in the genes and pathways identified may affect the role of the planum temporale in language cognition.

Abstract

In this paper we introduce Functional Data Analysis (FDA) as a tool for analyzing dynamic transitions in speech signals. FDA makes it possible to perform statistical analyses of sets of mathematical functions in the same way as classical multivariate analysis treats scalar measurement data. We illustrate the use of FDA with a reduction phenomenon affecting the French word c'était /setε/ 'it was', which can be reduced to [stε] in conversational speech. FDA reveals that the dynamics of the transition from [s] to [t] in fully reduced cases may still be different from the dynamics of [s] - [t] transitions in underlying /st/ clusters such as in the word stage.

Abstract

The study of phonetic contrasts and related phenomena, e.g. inter- and intra-speaker variability, often requires to analyse data in the form of measured time series, like f0 contours and formant trajectories. As a consequence, the investigator has to find suitable ways to reduce the raw and abundant numerical information contained in a bundle of time series into a small but sufficient set of numerical descriptors of their shape. This approach requires one to decide in advance which dynamic traits to include in the analysis and which not. For example, a rising pitch gesture may be represented by its duration and slope, hence reducing it to a straight segment, or by a richer coding specifying also whether (and how much) the rising contour is concave or convex, the latter being irrelevant in some context but crucial in others. Decisions become even more complex when a phenomenon is described by a multidimensional time series, e.g. by the first two formants. In this paper we introduce a methodology based on Functional Data Analysis (FDA) that allows the investigator to delegate most of the decisions involved in the quantitative description of multidimensional time series to the data themselves. FDA produces a data-driven parametrisation of the main shape traits present in the data that is visually interpretable, in the same way as slopes or peak heights are. These output parameters are numbers that are amenable to ordinary statistical analysis, e.g. linear (mixed effects) models. FDA is also able to capture correlations among different dimensions of a time series, e.g. between formants F1 and F2. We present FDA by means of an extended case study on diphthong – hiatus distinction in Spanish, a contrast that involves duration, formant trajectories and pitch contours.

Abstract

In order to make sense of the world, humans tend to see causation almost everywhere. Although most causal relations may seem straightforward, they are not always construed in the same way cross-culturally. In this study, we investigate concepts of ‘chance’, ‘coincidence’ or ‘randomness’ that refer to assumed relations between intention, action, and outcome in situations, and we ask how people from different cultures make sense of such non-law-like connections. Based on a framework proposed by Alicke (2000), we administered a task that aims to be a neutral tool for investigating causal construals cross-culturally and cross-linguistically. Members of four different cultural groups, rural Mayan Yucatec and Tseltal speakers from Mexico and urban students from Mexico and Germany, were presented with a set of scenarios involving various types of causal and non-causal relations and were asked to explain the described events. Three links varied as to whether they were present or not in the scenarios: Intention to Action, Action to Outcome, and Intention to Outcome. Our results show that causality is recognized in all four cultural groups. However, how causality and especially non-law-like causality are interpreted depends on the type of links, the cultural background and the language used. In all three groups, Action to Outcome is the decisive link for recognizing causality. Despite the fact that the two Mayan groups share similar cultural backgrounds, they display different ideologies regarding concepts of non-law causality. The data suggests that the concept of ‘chance’ is not universal, but seems to be an explanation that only some cultural groups draw on to make sense of specific situations. Of particular importance is the existence of linguistic concepts in each language that trigger ideas of causality in the responses from each cultural group

Abstract

The goal of this task is to investigate cross-cultural emotion categories in language and thought. This entry is designed to provide researchers with some guidelines to describe the emotional repertoire of a community from an emic perspective. The first objective is to offer ethnographic tools and a questionnaire in order to understand the semantics of emotional terms and the local conception of emotions. The second objective is to identify the local display rules of emotions in communicative interactions.

Abstract

Studies in relatives have generally yielded high heritability estimates for refractive error: twins 75–90%, families 15–70%. However, because related individuals often share a common environment, these estimates are inflated (via misallocation of unique/common environment variance). We calculated a lower-bound heritability estimate for refractive error free from such bias.
Between the ages 7 and 15 years, participants in the Avon Longitudinal Study of Parents and Children (ALSPAC) underwent non-cycloplegic autorefraction at regular research clinics. At each age, an estimate of the variance in refractive error explained by single nucleotide polymorphism (SNP) genetic variants was calculated using genome-wide complex trait analysis (GCTA) using high-density genome-wide SNP genotype information (minimum N at each age=3,404).
The variance in refractive error explained by the SNPs (“SNP heritability”) was stable over childhood: Across age 7–15 years, SNP heritability averaged 0.28 (SE=0.08, p<0.001). The genetic correlation for refractive error between visits varied from 0.77 to 1.00 (all p<0.001) demonstrating that a common set of SNPs was responsible for the genetic contribution to refractive error across this period of childhood. Simulations suggested lack of cycloplegia during autorefraction led to a small underestimation of SNP heritability (adjusted SNP heritability=0.35; SE=0.09). To put these results in context, the variance in refractive error explained (or predicted) by the time participants spent outdoors was <0.005 and by the time spent reading was <0.01, based on a parental questionnaire completed when the child was aged 8–9 years old.
Genetic variation captured by common SNPs explained approximately 35% of the variation in refractive error between unrelated subjects. This value sets an upper limit for predicting refractive error using existing SNP genotyping arrays, although higher-density genotyping in larger samples and inclusion of interaction effects is expected to raise this figure toward twin- and family-based heritability estimates. The same SNPs influenced refractive error across much of childhood. Notwithstanding the strong evidence of association between time outdoors and myopia, and time reading and myopia, less than 1% of the variance in myopia at age 15 was explained by crude measures of these two risk factors, indicating that their effects may be limited, at least when averaged over the whole population.

Abstract

Trumai, a genetically isolated language spoken in Brazil (Xingu reserve), is an example of an endangered language. Although the Trumai population consists of more than 100 individuals, only 51 people speak the language. The oral traditions are progressively dying. Given the current scenario, the documentation of this language and its cultural aspects is of great importance. In the framework of the DoBeS program (Documentation of Endangered Languages), the project "Documentation of Trumai" has selected and organized a collection of Trumai texts, with a multi-media representation of the corpus. Several kinds of information and data types are being included in the archive of the language: texts with audio and video recordings; written texts from educational materials; drawings; photos; songs; annotations in different formats; lexicon; field notes; results from scientific studies of the language (sound system, sketch grammar, comparative studies with other Xinguan languages), etc. All materials are integrated into the IMDI-Browser, a specialized tool for presenting and searching for linguistic data. This paper explores the processing phases and the results of the Trumai project taking into consideration the issue of how to combine the needs and wishes of field linguistics (content and research aspects) and the needs of archiving (structure and workflow aspects) in a well-organized corpus.

Abstract

There is growing evidence that addressees in interaction integrate the semantic information conveyed by speakers’ gestures. Little is known, however, about whether and how addressees’ attention to gestures and the integration of gestural information can be modulated. This study examines the influence of a social factor (speakers’ gaze to their own gestures), and two physical factors (the gesture’s location in gesture space and gestural holds) on addressees’ overt visual attention to gestures (direct fixations of gestures) and their uptake of gestural information. It also examines the relationship between gaze and uptake. The results indicate that addressees’ overt visual attention to gestures is affected both by speakers’ gaze and holds but for different reasons, whereas location in space plays no role. Addressees’ uptake of gesture information is only influenced by speakers’ gaze. There is little evidence of a direct relationship between addressees’ direct fixations of gestures and their uptake.

Abstract

This paper argues that speech-associated gestures can usefully inform studies exploring development of meaning in first and second language acquisition. The example domain is caused motion or placement meaning (putting a cup on a table) where acquisition problems have been observed and where adult native gesture use reflects crosslinguistically different placement verb semantics. Against this background, the paper summarises three studies examining the development of semantic representations in Dutch children acquiring Dutch, and adult learners’ acquiring Dutch and French placement verbs. Overall, gestures change systematically with semantic development both in children and adults and (1) reveal what semantic elements are included in current semantic representations, whether target-like or not, and (2) highlight developmental shifts in those representations. There is little evidence that gestures chiefly act as a support channel. Instead, the data support the theoretical notion that speech and gesture form an integrated system, opening new possibilities for studying the processes of acquisition.

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

This study examines to what extent English speakers of L2 Dutch reconstruct the meanings of placement verbs when moving from a general L1 verb of caused motion (put) to two specific caused posture verbs (zetten/leggen ‘set/lay’) in the L2 and whether the existence of low-frequency cognate forms in the L1 (set/lay) alleviates the reconstruction problem. Evidence from speech and gesture indicates that English speakers have difficulties with the specific verbs in L2 Dutch, initially looking for means to express general caused motion in L1-like fashion through over-generalisation. The gesture data further show that targetlike forms are often used to convey L1-like meaning. However, the differentiated use of zetten for vertical placement and dummy verbs (gaan ‘go’ and doen ‘do’) and intransitive posture verbs (zitten/staan/liggen ‘sit, stand, lie’) for horizontal placement, and a positive correlation between appropriate verb use and target-like gesturing suggest a beginning sensitivity to the semantic parameters of the L2 verbs and possible reconstruction.

Abstract

The aim of this chapter is to provide researchers with a tool kit of semi-experimental and experimental techniques for studying code-switching. It presents an overview of the current off-line and on-line research techniques, ranging from analyses of published bilingual texts of spontaneous conversations, to tightly controlled experiments. A multi-task approach used for studying code-switched sentence production in Papiamento-Dutch bilinguals is also exemplified.

Abstract

Gestures, the symbolic movements speakers perform while they speak, are systematically related to speech and language in non-trivial ways. This chapter presents an overview of what gestures can and cannot tell us about the monolingual and the bilingual mental lexicon. Gesture analysis opens for a broader view of the mental lexicon, targeting the interface between conceptual, semantic and syntactic aspects of event construal, and offers new possibilities for examining how languages co-exist and interact in bilinguals beyond the level of surface forms. The first section of this chapter gives a brief introduction to gesture studies and outlines the current views on the relationship between gesture, speech, and language. The second section targets the key questions for the study of the monolingual and bilingual lexicon, and illustrates the methods employed for addressing these questions. It further exemplifies systematic cross-linguistic patterns in gestural behaviour in monolingual and bilingual contexts. The final section discusses some implications of an expanded view of the multilingual lexicon that includes gesture, and outlines directions for future inquiry.

Abstract

A prototypical lexicon tool was implemented which was intended to allow researchers to collaboratively create lexicons of endangered languages. Increasingly often researchers documenting or analyzing a language work at different locations. Lexicons that evolve through continuous interaction between the collaborators can only be efficiently produced when it can be accessed and manipulated via the Internet. The SHAWEL tool was developed to address these needs; it makes use of a thin Java client and a central database solution.

Abstract

Analyses of gray matter concentration (GMC) deficits in patients with schizophrenia (Sz) have identified robust changes throughout the cortex. We assessed the relationships between diagnosis, overall symptom severity, and patterns of gray matter in the largest aggregated structural imaging dataset to date. We performed both source-based morphometry (SBM) and voxel-based morphometry (VBM) analyses on GMC images from 784 Sz and 936 controls (Ct) across 23 scanning sites in Europe and the United States. After correcting for age, gender, site, and diagnosis by site interactions, SBM analyses showed 9 patterns of diagnostic differences. They comprised separate cortical, subcortical, and cerebellar regions. Seven patterns showed greater GMC in Ct than Sz, while 2 (brainstem and cerebellum) showed greater GMC for Sz. The greatest GMC deficit was in a single pattern comprising regions in the superior temporal gyrus, inferior frontal gyrus, and medial frontal cortex, which replicated over analyses of data subsets. VBM analyses identified overall cortical GMC loss and one small cluster of increased GMC in Sz, which overlapped with the SBM brainstem component. We found no significant association between the component loadings and symptom severity in either analysis. This mega-analysis confirms that the commonly found GMC loss in Sz in the anterior temporal lobe, insula, and medial frontal lobe form a single, consistent spatial pattern even in such a diverse dataset. The separation of GMC loss into robust, repeatable spatial patterns across multiple datasets paves the way for the application of these methods to identify subtle genetic and clinical cohort effects.

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

This contribution focuses on the neural infrastructure for parsing and syntactic encoding. From an anatomical point of view, it is argued that Broca's area is an ill-conceived notion. Functionally, Broca's area and adjacent cortex (together Broca's complex) are relevant for language, but not exclusively for this domain of cognition. Its role can be characterized as providing the necessary infrastructure for unification (syntactic and semantic). A general proposal, but with required level of computational detail, is discussed to account for the distribution of labor between different components of the language network in the brain.Arguments are provided for the immediacy principle, which denies a privileged status for syntax in sentence processing. The temporal profile of event-related brain potential (ERP) is suggested to require predictive processing. Finally, since, next to speed, diversity is a hallmark of human languages, the language readiness of the brain might not depend on a universal, dedicated neural machinery for syntax, but rather on a shaping of the neural infrastructure of more general cognitive systems (e.g., memory, unification) in a direction that made it optimally suited for the purpose of communication through language.

Abstract

Language and communication are about the exchange of meaning. A key feature of understanding and producing language is the construction of complex meaning from more elementary semantic building blocks. The functional characteristics of this semantic unification process are revealed by studies using event related brain potentials. These studies have found that word meaning is assembled into compound meaning in not more than 500 ms. World knowledge, information about the speaker, co-occurring visual input and discourse all have an immediate impact on semantic unification, and trigger similar electrophysiological responses as sentence-internal semantic information. Neuroimaging studies show that a network of brain areas, including the left inferior frontal gyrus, the left superior/middle temporal cortex, the left inferior parietal cortex and, to a lesser extent their right hemisphere homologues are recruited to perform semantic unification.