Publications

Abstract

Clahsen's characterization of nondefault inflection as based exclusively on lexical entries does not capture the full range of empirical data on German inflection. In the verb system differential effects of lexical frequency seem to be input-related rather than affecting morphological production. In the noun system, the generalization properties of -n and -e plurals exceed mere analogy-based productivity.

Abstract

In order to separate the cognitive processes associated with phonological encoding and the use of a visual word form lexicon in reading, it is desirable to compare the processing of words presented in a visually familiar form with words in a visually unfamiliar form. Japanese Kana orthography offers this possibility. Two phonologically equivalent but visually dissimilar syllabaries allow the writing of, for example, foreign loanwords in two ways, only one of which is visually familiar. Familiarly written words, unfamiliarly written words, and pseudowords were presented in both Kana syllabaries (yielding six conditions in total) to participants during an fMRI measurement with a silent articulation task (Experiment 1) and a phonological lexical decision task (Experiment 2) using an event-related design. Consistent over two experimental tasks, the three different stimulus types (familiar, unfamiliar, and pseudoword) were found to activate selectively different brain regions previously associated with phonological encoding and word retrieval or meaning. Compatible with the predictions of the dual-route model for reading, pseudowords and visually unfamiliar words, which have to be read using phonological assembly, caused an increase in brain activity in left inferior frontal regions (BA 44/47), as compared to visually familiar words. Visually familiar and unfamiliar words were found to activate a range of areas associated with lexico-semantic processing more strongly than pseudowords, such as the left and right temporo-parietal region (BA 39/40), a region in the left middle/inferior temporal gyrus (BA 20/21), and the posterior cingulate (BA 31).

Abstract

Bioacoustics increasingly relies on large datasets and computational methods. The need to batch-process large amounts of data and the increased focus on algorithmic processing require software tools. To optimally assist in a bioacoustician’s workflow, software tools need to be as simple and effective as possible. Five years ago, the Python package Parselmouth was released to provide easy and intuitive access to all functionality in the Praat software. Whereas Praat is principally designed for phonetics and speech processing, plenty of bioacoustics studies have used its advanced acoustic algorithms. Here, we evaluate existing usage of Parselmouth and discuss in detail several studies which used the software library. We argue that Parselmouth has the potential to be used even more in bioacoustics research, and suggest future directions to be pursued with the help of Parselmouth.

Abstract

Natural fast speech differs from normal-rate speech with respect to its temporal pattern. Previous results showed that word intelligibility of heavily artificially time-compressed speech could not be improved by making its temporal pattern more similar to that of natural fast speech. This might have been due to the extrapolation of timing rules for natural fast speech to rates that are much faster than can be attained by human speakers. The present study investigates whether, at a speech rate that human speakers can attain, artificially time-compressed speech is easier to process if its timing pattern is similar to that of naturally produced fast speech. Our first experiment suggests, however, that word processing speed was slowed down, relative to linear compression. In a second experiment, word processing of artificially time-compressed speech was compared with processing of naturally produced fast speech. Even when naturally produced fast speech is perfectly intelligible, its less careful articulation, combined with the changed timing pattern, slows down processing, relative to linearly time-compressed speech. Furthermore, listeners preferred artificially time-compressed speech over naturally produced fast speech. These results suggest that linearly time-compressed speech has both a temporal and a segmental advantage over natural fast speech.

Abstract

This study investigated the encoding of syllable boundary information during speech production in Dutch. Based on Levelt's model of phonological encoding, we hypothesized segments and syllable boundaries to be encoded in an incremental way. In a selfmonitoring experiment, decisions about the syllable affiliation (first or second syllable) of a pre-specified consonant, which was the third phoneme in a word, were required (e.g., ka.No canoe vs. kaN.sel pulpit ; capital letters indicate pivotal consonants, dots mark syllable boundaries). First syllable responses were faster than second syllable responses, indicating the incremental nature of segmental encoding and syllabification during speech production planning. The results of the experiment are discussed in the context of Levelt 's model of phonological encoding.

Abstract

Three experiments are reported that examined whether stem complexity plays a role in inflecting polymorphemic words in language production. Experiment 1 showed that preparation effects for words with polymorphemic stems are larger when they are produced among words with constant inflectional structures compared to words with variable inflectional structures and simple stems. This replicates earlier findings for words with monomorphemic stems (Janssen et al., 2002). Experiments 2 and 3 showed that when inflectional structure is held constant, the preparation effects are equally large with simple and compound stems, and with compound and complex adjectival stems. These results indicate that inflectional encoding is blind to the complexity of the stem, which suggests that specific inflectional rather than generic morphological frames guide the generation of inflected forms in speaking words.

Abstract

As people find their way through their environment, objects at navigationally relevant locations can serve as crucial landmarks. The parahippocampal gyrus has previously been shown to be involved in object and scene recognition. In the present study, we investigated the neural representation of navigationally relevant locations. Healthy human adults viewed a route through a virtual museum with objects placed at intersections (decision points) or at simple turns (non-decision points). Event-related functional magnetic resonance imaging (fMRI) data were acquired during subsequent recognition of the objects in isolation. Neural activity in the parahippocampal gyrus reflected the navigational relevance of an object's location in the museum. Parahippocampal responses were selectively increased for objects that occurred at decision points, independent of attentional demands. This increase occurred for forgotten as well as remembered objects, showing implicit retrieval of navigational information. The automatic storage of relevant object location in the parahippocampal gyrus provides a part of the neural mechanism underlying successful navigation.

Abstract

In 7 experiments the authors investigated the locus of word frequency effects in speech production. Experiment 1 demonstrated a frequency effect in picture naming that was robust over repetitions. Experiments 2, 3, and 7 excluded contributions from object identification and initiation of articulation. Experiments 4 and 5 investigated whether the effect arises in accessing the syntactic word (lemma) by using a grammatical gender decision task. Although a frequency effect was found, it dissipated under repeated access to word's gender. Experiment 6 tested whether the robust frequency effect arises in accessing the phonological form (lexeme) by having Ss translate words that produced homophones. Low-frequent homophones behaved like high-frequent controls, inheriting the accessing speed of their high-frequent homophone twins. Because homophones share the lexeme, not the lemma, this suggests a lexeme-level origin of the robust effect.

Abstract

This study investigates the time-course of information processing in both visual as well as in the auditory speech as used for word identification in face-to-face communication. It extends the limited previous research on this topic and provides a valuable database for future research in audiovisual speech perception. An evaluation of models of speech perception by ear and eye in their ability to account for the audiovisual gating data shows a superior role of the fuzzy logical model of perception (FLMP) [1] over additive models of perception. A new dynamic version of the FLMP seems to be a promising model to account for the complex interplay of perceptual and cognitive information in audiovisual spoken word recognition.

Abstract

In face-to-face communication, information from the face as well as from the voice contributes to the identification of spoken words. This dissertation investigates the time-course of the evaluation and integration of visual and auditory speech in audiovisual word identification. A large-scale audiovisual gating study extends previous research on this topic by (1) using a set of words that includes all possible initial consonants in English in three vowel contexts, (2) tracking the information processing for individual words not only across modalities, but also over time, and (3) testing quantitative models of the time-course of multimodal word recognition. There was an advantage in accuracy for audiovisual speech over auditory-only and visual-only speech. Auditory performance was, however, close to ceiling while performance on visual-only trials was near the floor of the scale, but well above chance. Visual information was used at all gates to identify the presented words. Information theoretic feature analyses of the confusion matrices revealed that the auditory signal is highly informative about voicing, manner, frication, duration, and place of articulation. Visual speech is mostly informative about place of articulation, but also about frication and duration. The auditory signal provides more information about the place of articulation for back consonants, whereas the visual signal provides more information for the labial consonants. The data were sufficient to discriminate between models of audiovisual word recognition. The Fuzzy Logical Model of Perception (FLMP; Massaro, 1998) gave a better account of the confusion matrix data than additive models of perception. A dynamic version of the FLMP was expanded to account for the evaluation and integration of information over time. This dynamic FLMP provided a better description of the data than dynamic additive competitor models. The present study builds a good foundation to investigate the role of the complex interplay between stimulus information and the structure of the lexicon. It provides an important step in building a formal representation of a lexical dynamic FLMP that can account not only for the time-course of speech information and its perceptual processing, but also for lexical influences.

Abstract

Mutations of the epidermal growth factor receptor (EGFR) gene are found at a relatively high frequency in glioma, with the most common being the de2-7 EGFR (or EGFRvIII). This mutation arises from an in-frame deletion of exons 2–7, which removes 267 amino acids from the extracellular domain of the receptor. Despite being unable to bind ligand, the de2-7 EGFR is constitutively active at a low level. Transfection of human glioma cells with the de2-7 EGFR has little effect in vitro, but when grown as tumor xenografts this mutated receptor imparts a dramatic growth advantage. We have now mapped the phosphorylation pattern of de2-7 EGFR, both in vivo and in vitro, using a panel of antibodies unique to the different phosphorylated tyrosine residues. Phosphorylation of de2-7 EGFR was detected constitutively at all tyrosine sites surveyed both in vitro and in vivo, including tyrosine 845, a known target in the wild-type EGFR for src kinase. There was a substantial upregulation of phosphorylation at every tyrosine residue of the de2-7 EGFR when cells were grown in vivo compared to the receptor isolated from cells cultured in vitro. Upregulation of phosphorylation could be mimicked in vitro by the addition of specifi c components of the ECM such as collagen via an integrin-dependent mechanism. Since this increase in in vivo phosphorylation enhances de2-7 EGFR signaling, this observation explains why the growth enhancement mediated by de2-7 EGFR is largely restricted to the in vivo environment. In a second set of experiments we analyzed the interaction between EGFRvIII and ErbB2. Co-expression of these proteins in NR6 cells, a mouse fi broblast line devoid of ErbB family members, dramatically enhanced in vivo tumorigenicity of these cells compared to cells expressing either protein alone. Detailed analysis of these xenografts demonstrated that EGFRvIII could heterodimerize and transphosphorylate the ErbB2. Since both EGFRvIII and ErbB2 are commonly expressed at gliomas, this data suggests that the co-expression of these two proteins may enhance glioma tumorigenicity.

Abstract

Mutations of the epidermal growth factor receptor (EGFR) gene are found at a relatively high frequency in glioma, with the most common being the de2-7 EGFR (or EGFRvIII). This mutation arises from an in-frame deletion of exons 2-7, which removes 267 amino acids from the extracellular domain of the receptor. Despite being unable to bind ligand, the de2-7 EGFR is constitutively active at a low level. Transfection of human glioma cells with the de2-7 EGFR has little effect in vitro, but when grown as tumor xenografts this mutated receptor imparts a dramatic growth advantage. We mapped the phosphorylation pattern of de2-7 EGFR, both in vivo and in vitro, using a panel of antibodies specific for different phosphorylated tyrosine residues. Phosphorylation of de2-7 EGFR was detected constitutively at all tyrosine sites surveyed in vitro and in vivo, including tyrosine 845, a known target in the wild-type EGFR for src kinase. There was a substantial upregulation of phosphorylation at every yrosine residue of the de2-7 EGFR when cells were grown in vivo compared to the receptor isolated from cells cultured in vitro. Upregulation of phosphorylation at tyrosine 845 could be stimulated in vitro by the addition of specific components of the ECM via an integrindependent mechanism. These observations may partially explain why the growth enhancement mediated by de2-7 EGFR is largely restricted to the in vivo environment

Abstract

Retaining detailed representations of unstressed syllables is a logical prerequisite for infants' use of probabilistic phonotactics to segment iambic words from fluent speech. The head-turn preference study was used to investigate the nature of English- learners' representations of iambic word onsets. Fifty-four 10.5-month-olds were familiarized to passages containing the nonsense iambic word forms ginome and tupong. Following familiarization, infants were either tested on familiar (ginome and tupong) or near-familiar (pinome and bupong) versus unfamiliar (kidar and mafoos) words. Infants in the familiar test group (familiar vs. unfamiliar) oriented significantly longer to familiar than unfamiliar test items, whereas infants in the near-familiar test group (near-familiar vs. unfamiliar) oriented equally long to near-familiar and unfamiliar test items. Our results provide evidence that infants retain fairly detailed representations of unstressed syllables and therefore support the hypothesis that infants use phonotactic cues to find words in fluent speech.

Abstract

In early Dutch learner varieties, there is no evidence of finiteness being a functional category. There is no V2nd: no correlation between inflectional morphology and movement. Initially, learners express the illocutive function of finiteness through the use of illocutive markers, with the non-use of an illocutive marker expressing the default illocutive function of assertion. Illocutive markers are functioning as adjuncts with scope over the predicate. Illocutive markers become re-analysed as functional elements.The driving force is the acquisition of the auxiliary verbs that occur with past participles. It leads to a reanalysis of illocutive markers as two separate elements: an auxiliary verb and a scope adverb. The (modal) auxiliary carries illocutive function. Lexical verb-argument structure (including the external argument) occurs within the domain of the auxiliary verb. The predicate as the focus constituent occurs within the domain of a scope adverb. This reanalysis establishes a position for the external argument within the domain of AUX. The acquisition of AUX causes the acquisition of a (hierarchical) structure with a complement as a constituent which represents an underlying verb-argument structure, a predicate as the domain of elements that are in focus, and an external (specifier) position as a landing site for elements with topic function.

Abstract

Unilateral anterior disc displacement (uADD) has been shown to affect the contralateral joints qualitatively. This study aims to assess the quantitative T2 values of the articular disc and retrodiscal tissue of patients with uADD at 1.5 Tesla (T). The study included 65 uADD patients and 17 volunteers. The regions of interest on T2 maps were evaluated. The affected joints demonstrated significantly higher articular disc T2 values (31.5 ± 3.8 ms) than those of the unaffected joints (28.9 ± 4.5 ms) (P < 0.001). For retrodiscal tissue, T2 values of the unaffected (37.8 ± 5.8 ms) and affected joints (41.6 ± 7.1 ms) were significantly longer than those of normal volunteers (34.4 ± 3.2 ms) (P < 0.001). Furthermore, uADD without reduction (WOR) joints (43.3 ± 6.8 ms) showed statistically higher T2 values than the unaffected joints of both uADD with reduction (WR) (33.9 ± 3.8 ms) and uADDWOR (38.9 ± 5.8 ms), and the affected joints of uADDWR (35.8 ± 4.4 ms). The mean T2 value of the unaffected joints of uADDWOR was significantly longer than that of healthy volunteers (P < 0.001). These results provided quantitative evidence for the influence of the affected joints on the contralateral joints.

Abstract

We investigated prediction skills of adult heritage speakers and the role of written and spoken language experience on predictive processing. Using visual world eye-tracking, we focused on predictive use of case-marking cues in verb-medial and verb-final sentences in Turkish with adult Turkish heritage speakers (N = 25) and Turkish monolingual speakers (N = 24). Heritage speakers predicted in verb-medial sentences (when verb-semantic and case-marking cues were available), but not in verb-final sentences (when only case-marking cues were available) while monolinguals predicted in both. Prediction skills of heritage speakers were modulated by their spoken language experience in Turkish and written language experience in both languages. Overall, these results strongly suggest that verb-semantic information is needed to scaffold the use of morphosyntactic cues for prediction in heritage speakers. The findings also support the notion that both spoken and written language experience play an important role in predictive spoken language processing.

Abstract

When learning a spoken second language (L2), words overlapping in form and meaning with one’s native language (L1) help break into the new language. When non-signing speakers learn a sign language as L2, such forms are absent because of the modality differences (L1:speech, L2:sign). In such cases, non-signing speakers might use iconic form-meaning mappings in signs or their own gestural experience as gateways into the to-be-acquired sign language. Here, we investigated how both these factors may contribute jointly to the acquisition of sign language vocabulary by hearing non-signers. Participants were presented with three types of sign in NGT (Sign Language of the Netherlands): arbitrary signs, iconic signs with high or low gesture overlap. Signs that were both iconic and highly overlapping with gestures boosted learning most at first exposure, and this effect remained the day after. Findings highlight the influence of modality-specific factors supporting the acquisition of a signed lexicon.

Abstract

The 8p23.1 duplication syndrome is a rare genetic condition with an estimated prevalence rate of 1 out of 58,000. Although the syndrome was associated with speech and language delays, a comprehensive assessment of speech and language functions has not been undertaken in this population. To address this issue, the present study reports rigorous speech and language, in addition to oral-facial and developmental, assessment of a 50-month-old Turkish-speaking boy who was diagnosed with the 8p23.1 duplication syndrome. Standardized tests of development, articulation and phonology, receptive and expressive language and a language sample analysis were administered to characterize speech and language skills in the patient. The language sample was obtained in an ecologically valid, free play and conversation context. The language sample was then analyzed and compared to a database of age-matched typically-developing children (n = 33) in terms of intelligibility, morphosyntax, semantics/vocabulary, discourse, verbal facility and percentage of errors at word and utterance levels. The results revealed mild to severe problems in articulation and phonology, receptive and expressive language skills, and morphosyntax (mean length of utterance in morphemes). Future research with larger sample sizes and employing detailed speech and language assessment is needed to delineate the speech and language profile in individuals with the 8p23.1 duplication syndrome, which will guide targeted speech and language interventions.

Abstract

We outline an algorithm capable of generating varied but natural sounding sequences of argument NPs in subordinate clauses of German, a semi-free word order language. In order to attain the right level of output flexibility, the algorithm considers (1) the relevant lexical properties of the head verb (not only transitivity type but also reflexivity, thematic relations expressed by the NPs, etc.), and (2) the animacy and definiteness values of the arguments, and their length. The relevant statistical data were extracted from the NEGRA–II treebank and from hand-coded features for animacy and definiteness. The algorithm maps the relevant properties onto “primary” versus “secondary” placement options in the generator. The algorithm is restricted in that it does not take into account linear order determinants related to the sentence’s information structure and its discourse context (e.g. contrastiveness). These factors may modulate the above preferences or license “tertiary” linear orders beyond the primary and secondary options considered here.

Abstract

Listeners cannot recognize highly reduced word forms in isolation, but they can do so when these forms are presented in context (Ernestus, Baayen, & Schreuder, 2002). This suggests that not all possible surface forms of words have equal status in the mental lexicon. The present study shows that the reduced forms are linked to the canonical representations in the mental lexicon, and that these latter representations induce reconstruction processes. Listeners restore suffixes that are partly or completely missing in reduced word forms. A series of phoneme-monitoring experiments reveals the nature of this restoration: the basis for suffix restoration is mainly phonological in nature, but orthography has an influence as well.

Abstract

Previous work has shown that Dutch listeners use prosodic information in the speech signal to optimise morphological processing: Listeners are sensitive to prosodic differences between a noun stem realised in isolation and a noun stem realised as part of a plural form (in which the stem is followed by an unstressed syllable). The present study, employing a lexical decision task, provides an additional demonstration of listeners' sensitivity to prosodic cues in the stem. This sensitivity is shown for two languages that differ in morphological productivity: Dutch and English. The degree of morphological productivity does not correlate with listeners' sensitivity to prosodic cues in the stem, but it is reflected in differential sensitivities to the word-specific log odds ratio of encountering an unshortened stem (i.e., a stem in isolation) versus encountering a shortened stem (i.e., a stem followed by a suffix consisting of one or more unstressed syllables). In addition to being sensitive to the prosodic cues themselves, listeners are also sensitive to the probabilities of occurrence of these prosodic cues.

Abstract

It has recently been shown that listeners use systematic differences in vowel length and intonation to resolve ambiguities between onset-matched simple words (Davis, Marslen-Wilson, & Gaskell, 2002; Salverda, Dahan, & McQueen, 2003). The present study shows that listeners also use prosodic information in the speech signal to optimize morphological processing. The precise acoustic realization of the stem provides crucial information to the listener about the morphological context in which the stem appears and attenuates the competition between stored inflectional variants. We argue that listeners are able to make use of prosodic information, even though the speech signal is highly variable within and between speakers, by virtue of the relative invariance of the duration of the onset. This provides listeners with a baseline against which the durational cues in a vowel and a coda can be evaluated. Furthermore, our experiments provide evidence for item-specific prosodic effects.

Abstract

This dissertation investigates two seemingly contradictory properties of the speech perception system. On the one hand, listeners are extremely sensitive to the fine phonetic details in the speech signal. These subtle acoustic cues can reduce the temporal ambiguity between words that show initial segmental overlap, and can guide lexical activation. On the other hand, comprehension does not seem to be hampered at all by the drastic reductions that typically occur in casual speech. Complete segments, and sometimes even complete syllables, may be missing, but comprehension is seemingly unaffected. This thesis aims at elucidating how words are represented and accessed in the mental lexicon, by investigating these contradictory phenomena for the domain of morphology

Abstract

In Dutch, high-frequency words with the suffix -lijk are often highly reduced in spontaneous unscripted speech. This study addressed socio-geographic variation in the reduction of such words against the backdrop of the variation in their use in written and spoken Dutch. Multivariate analyses of the frequencies with which the words were used in a factorially contrasted set of subcorpora revealed signi ficant variation involving the speaker's country, sex, and education level for spoken Dutch, and involving country and register for written Dutch. Acoustic analyses revealed that Dutch men reduced most often, while Flemish highly educated women reduced least. Two linguistic context effects emerged, one prosodic, and the other pertaining to the flow of information. Words in sentence final position showed less reduction, while words that were better predictable from the preceding word in the sentence(based on mutual information) tended to be reduced more often. The increased probability of reduction for forms that are more predictable in context, combined with the loss of the suffix in the more extremely reduced forms, suggests that highfrequency words in -lijk are undergoing a process of erosion that causes them to gravitate towards monomorphemic function words.

Abstract

Drozd's critique of Crain & Thornton's (C&T) (1998) book Investigations in Universal Grammar (IUG) raises many issues concerning theory and experimental design within generative approaches to language acquisition. I focus here on one of the strongest theoretical claims of the Modularity Matching Model (MMM): continuity of processing. For reasons different to Drozd, I argue that the assumption is tenuous. Furthermore, I argue that the focus of the MMM and the methodological prescriptions contained in IUG are too narrow to capture language acquisition.

Abstract

This paper reports on an investigation of children's (aged 3;5–9;8) comprehension of sentences containing ambiguity of prepositional phrase (PP) attachment. Results from a picture selection study (N=90) showed that children use verb semantics and preposition type to resolve the ambiguity, with older children also showing sensitivity to the definiteness of the object NP as a cue to interpretation. Study 2 investigated three- and five-year-old children's (N=47) ability to override an instrumental interpretation of ambiguous PPs in order to process attributes of the referential scene. The results showed that while five-year-olds are capable of incorporating aspects of the referential scene into their interpretations, three-year-olds are not as successful. Overall, the results suggest that children are attuned very early to the lexico-semantic co-occurrences that have been shown to aid ambiguity resolution in adults, but that more diffuse cues to interpretation are used only later in development

Abstract

Somatic practices frequently use imagery, typically via verbal instructions, to scaffold sensorimotor organization and experience, a phenomenon we term “introjection”. We argue that introjection is an imagery practice in which sensorimotor and conceptual aspects are co-orchestrated, suggesting the necessity of crosstalk between somatics, phenomenology, psychology, embodied-enactive cognition, and linguistic research on embodied simulation. We presently focus on the scarcely addressed details of the process necessary to enact instructions of a literal or metaphoric nature through the body. Based on vignettes from dance, Feldenkrais, and Taichi practice, we describe introjection as a complex form of processual sense-making, in which context-interpretive, mental, attentional and physical sub-processes recursively braid. Our analysis focuses on how mental and body-related processes progressively align, inform and augment each other. This dialectic requires emphasis on the active body, which implies that uni-directional models (concept ⇒ body) are inadequate and should be replaced by interactionist alternatives (concept ⇔ body). Furthermore, we emphasize that both the source image itself and the body are specifically conceptualized for the context through constructive operations, and both evolve through their interplay. At this level introjection employs representational operations that are embedded in enactive dynamics of a fully situated person.

Abstract

Pauses during continuous speech, particularly those that occur within clauses, are thought to reflect the planning of forthcoming verbal output. We used functional Magnetic Resonance Imaging (fMRI) to examine their neural correlates. Six volunteers were scanned while describing seven Rorschach inkblots, producing 3 min of speech per inkblot. In an event-related design, the level of blood oxygenation level dependent (BOLD) contrast during brief speech pauses (mean duration 1.3 s, SD 0.3 s) during overt speech was contrasted with that during intervening periods of articulation. We then examined activity associated with pauses that occurred within clauses and pauses that occurred between grammatical junctions. Relative to articulation during speech, pauses were associated with activation in the banks of the left superior temporal sulcus (BA 39/22), at the temporoparietal junction. Continuous speech was associated with greater activation bilaterally in the inferior frontal (BA 44/45), middle frontal (BA 8) and anterior cingulate (BA 24) gyri, the middle temporal sulcus (BA 21/22), the occipital cortex and the cerebellum. Left temporal activation was evident during pauses that occurred within clauses but not during pauses at grammatical junctions. In summary, articulation during continuous speech involved frontal, temporal and cerebellar areas, while pausing was associated with activity in the left temporal cortex, especially when this occurred within a clause. The latter finding is consistent with evidence that within-clause pauses are a correlate of speech planning and in particular lexical retrieval.

Abstract

The size of the human head is highly heritable, but genetic drivers of its variation within the general population remain unmapped. We perform a genome-wide association study on head size (N = 80,890) and identify 67 genetic loci, of which 50 are novel. Neuroimaging studies show that 17 variants affect specific brain areas, but most have widespread effects. Gene set enrichment is observed for various cancers and the p53, Wnt, and ErbB signaling pathways. Genes harboring lead variants are enriched for macrocephaly syndrome genes (37-fold) and high-fidelity cancer genes (9-fold), which is not seen for human height variants. Head size variants are also near genes preferentially expressed in intermediate progenitor cells, neural cells linked to evolutionary brain expansion. Our results indicate that genes regulating early brain and cranial growth incline to neoplasia later in life, irrespective of height. This warrants investigation of clinical implications of the link between head size and cancer.