Publications

Abstract

It is widely accepted that specific memory processes, such as serial-order memory, are involved in written language development and predictive of reading and spelling abilities. The reverse question, namely whether orthographic abilities also affect serial-order memory, has hardly been investigated. In the current study, we compared 20 illiterate people with a group of 20 literate matched controls on a verbal and a visuospatial version of the Hebb paradigm, measuring both short- and long-term serial-order memory abilities. We observed better short-term serial-recall performance for the literate compared with the illiterate people. This effect was stronger in the verbal than in the visuospatial modality, suggesting that the improved capacity of the literate group is a consequence of learning orthographic skills. The long-term consolidation of ordered information was comparable across groups, for both stimulus modalities. The implications of these findings for current views regarding the bi-directional interactions between memory and written language development are discussed.

Abstract

Spinocerebellar ataxia type 23 is caused by mutations in PDYN, which encodes the opioid neuropeptide precursor protein, prodynorphin. Prodynorphin is processed into the opioid peptides, α-neoendorphin, and dynorphins A and B, that normally exhibit opioid-receptor mediated actions in pain signalling and addiction. Dynorphin A is likely a mutational hotspot for spinocerebellar ataxia type 23 mutations, and in vitro data suggested that dynorphin A mutations lead to persistently elevated mutant peptide levels that are cytotoxic and may thus play a crucial role in the pathogenesis of spinocerebellar ataxia type 23. To further test this and study spinocerebellar ataxia type 23 in more detail, we generated a mouse carrying the spinocerebellar ataxia type 23 mutation R212W in PDYN. Analysis of peptide levels using a radioimmunoassay shows that these PDYNR212W mice display markedly elevated levels of mutant dynorphin A, which are associated with climber fibre retraction and Purkinje cell loss, visualized with immunohistochemical stainings. The PDYNR212W mice reproduced many of the clinical features of spinocerebellar ataxia type 23, with gait deficits starting at 3 months of age revealed by footprint pattern analysis, and progressive loss of motor coordination and balance at the age of 12 months demonstrated by declining performances on the accelerating Rotarod. The pathologically elevated mutant dynorphin A levels in the cerebellum coincided with transcriptionally dysregulated ionotropic and metabotropic glutamate receptors and glutamate transporters, and altered neuronal excitability. In conclusion, the PDYNR212W mouse is the first animal model of spinocerebellar ataxia type 23 and our work indicates that the elevated mutant dynorphin A peptide levels are likely responsible for the initiation and progression of the disease, affecting glutamatergic signalling, neuronal excitability, and motor performance. Our novel mouse model defines a critical role for opioid neuropeptides in spinocerebellar ataxia, and suggests that restoring the elevated mutant neuropeptide levels can be explored as a therapeutic intervention.

Abstract

Blindsight refers to the observation of residual visual abilities in the hemianopic field of patients without a functional V1. Given the within- and between-subject variability in the preserved abilities and the phenomenal experience of blindsight patients, the fine-grained description of the phenomenon is still debated. Here we tested a patient with established “perceptual” and “attentional” blindsight (c.f. Danckert and Rossetti, 2005). Using a pointing paradigm patient MS, who suffers from a complete left homonymous hemianopia, showed clear above chance manual localisation of ‘unseen’ targets. In addition, target presentations in his blind field led MS, on occasion, to spontaneous responses towards his sighted field. Structural and functional magnetic resonance imaging was conducted to evaluate the magnitude of V1 damage. Results revealed the presence of a calcarine sulcus in both hemispheres, yet his right V1 is reduced, structurally disconnected and shows no fMRI response to visual stimuli. Thus, visual stimulation of his blind field can lead to “action blindsight” and spontaneous antipointing, in absence of a functional right V1. With respect to the antipointing, we suggest that MS may have registered the stimulation and subsequently presumes it must have been in his intact half field.

Abstract

Previous research [9] suggests that Dutch learners of (British) English are not able to express sarcasm prosodically in their L2. The present study investigates whether explicit training on the prosodic markers of sarcasm in English can improve learners’ realisation of sarcasm. Sarcastic speech was elicited in short simulated telephone conversations between Dutch advanced learners of English and a native British English-speaking ‘friend’ in two sessions, fourteen days apart. Between the two sessions, participants were trained by means of (1) a presentation, (2) directed independent practice, and (3) evaluation of participants’ production and individual feedback in small groups. L1 British English-speaking raters subsequently evaluated the degree of sarcastic sounding in the participants’ responses on a five-point scale. It was found that significantly higher sarcasm ratings were given to L2 learners’ production obtained after the training than that obtained before the training; explicit training on prosody has a positive effect on learners’ production of sarcasm.

Abstract

Sentence comprehension requires the retrieval of single word information from long-term memory, and the integration of this information into multiword representations. The current functional magnetic resonance imaging study explored the hypothesis that the left posterior temporal gyrus supports the retrieval of lexical-syntactic information, whereas left inferior frontal gyrus (LIFG) contributes to syntactic unification. Twenty-eight subjects read sentences and word sequences containing word-category (noun–verb) ambiguous words at critical positions. Regions contributing to the syntactic unification process should show enhanced activation for sentences compared to words, and only within sentences display a larger signal for ambiguous than unambiguous conditions. The posterior LIFG showed exactly this predicted pattern, confirming our hypothesis that LIFG contributes to syntactic unification. The left posterior middle temporal gyrus was activated more for ambiguous than unambiguous conditions (main effect over both sentences and word sequences), as predicted for regions subserving the retrieval of lexical-syntactic information from memory. We conclude that understanding language involves the dynamic interplay between left inferior frontal and left posterior temporal regions.

Abstract

POU3F3, also referred to as Brain-1, is a well-known transcription factor involved in the development of the central nervous system, but it has not previously been associated with a neurodevelopmental disorder. Here, we report the identification of 19 individuals with heterozygous POU3F3 disruptions, most of which are de novo variants. All individuals had developmental delays and/or intellectual disability and impairments in speech and language skills. Thirteen individuals had characteristic low-set, prominent, and/or cupped ears. Brain abnormalities were observed in seven of eleven MRI reports. POU3F3 is an intronless gene, insensitive to nonsense-mediated decay, and 13 individuals carried protein-truncating variants. All truncating variants that we tested in cellular models led to aberrant subcellular localization of the encoded protein. Luciferase assays demonstrated negative effects of these alleles on transcriptional activation of a reporter with a FOXP2-derived binding motif. In addition to the loss-of-function variants, five individuals had missense variants that clustered at specific positions within the functional domains, and one small in-frame deletion was identified. Two missense variants showed reduced transactivation capacity in our assays, whereas one variant displayed gain-of-function effects, suggesting a distinct pathophysiological mechanism. In bioluminescence resonance energy transfer (BRET) interaction assays, all the truncated POU3F3 versions that we tested had significantly impaired dimerization capacities, whereas all missense variants showed unaffected dimerization with wild-type POU3F3. Taken together, our identification and functional cell-based analyses of pathogenic variants in POU3F3, coupled with a clinical characterization, implicate disruptions of this gene in a characteristic neurodevelopmental disorder.

Abstract

The anterior cingulate cortex (ACC) and the dorsolateral prefrontal cortex (DLPFC) are involved in conflict detection and
conflict resolution, respectively. Here, we investigate how lifelong bilingualism induces neuroplasticity to these structures by
employing a novel analysis of behavioural performance. We correlated grey matter volume (GMV) in seniors reported by
Abutalebi et al. (2015) with behavioral Flanker task performance fitted using the diffusion model (Ratcliff, 1978). As
predicted, we observed significant correlations between GMV in the DLPFC and Flanker performance. However, for
monolinguals the non-decision time parameter was significantly correlated with GMV in the left DLPFC, whereas for
bilinguals the correlation was significant in the right DLPFC. We also found a significant correlation between age and GMV
in left DLPFC and the non-decision time parameter for the conflict effect for monolinguals only.
We submit that this is due to cumulative demands on cognitive control over a lifetime of bilingual language processing

Abstract

In language comprehension, a variety of contextual cues act in unison to render upcoming words more or less predictable. As a sentence unfolds, we use prior context (sentential constraints) to predict what the next words might be. Additionally, in a conversation, we can predict upcoming sounds through observing the mouth movements of a speaker (visual constraints). In electrophysiological studies, effects of visual constraints have typically been observed early in language processing, while effects of sentential constraints have typically been observed later. We hypothesized that the visual and the sentential constraints might feed into the same predictive process such that effects of sentential constraints might also be detectable early in language processing through modulations of the early effects of visual salience. We presented participants with audiovisual speech while recording their brain activity with magnetoencephalography. Participants saw videos of a person saying sentences where the last word was either sententially constrained or not, and began with a salient or non-salient mouth movement. We found that sentential constraints indeed exerted an early (N1) influence on language processing. Sentential modulations of the N1 visual predictability effect were visible in brain areas associated with semantic processing, and were differently expressed in the two hemispheres. In the left hemisphere, visual and sentential constraints jointly suppressed the auditory evoked field, while the right hemisphere was sensitive to visual constraints only in the absence of strong sentential constraints. These results suggest that sentential and visual constraints can jointly influence even very early stages of audiovisual speech comprehension.

Abstract

Humans have a strong proclivity for structuring and patterning stimuli: Whether in space or time, we tend to mentally order stimuli in our environment and organize them into units with specific types of relationships. A crucial prerequisite for such organization is the cognitive ability to discern and process regularities among multiple stimuli. To investigate the evolutionary roots of this cognitive capacity, we tested chimpanzees—which, along with bonobos, are our closest living relatives—for simple, variable distance dependency processing in visual patterns. We trained chimpanzees to identify pairs of shapes either linked by an arbitrary learned association (arbitrary associative dependency) or a shared feature (same shape, feature-based dependency), and to recognize strings where items related to either of these ways occupied the first (leftmost) and the last (rightmost) item of the stimulus. We then probed the degree to which subjects generalized this pattern to new colors, shapes, and numbers of interspersed items. We found that chimpanzees can learn and generalize both types of dependency rules, indicating that the ability to encode both feature-based and arbitrary associative regularities over variable distances in the visual domain is not a human prerogative. Our results strongly suggest that these core components of human structural processing were already present in our last common ancestor with chimpanzees.

Abstract

Flexibly adapting social behavior to social and environmental challenges helps to alleviate glucocorticoid (GC) levels, which may have positive fitness implications for an individual. For primates, the predominant social behavior is grooming. Giving grooming to others is particularly efficient in terms of GC mitigation. However, grooming is confined by certain limitations such as time constraints or restricted access to other group members. For instance, dominance hierarchies may impact grooming partner availability in primate societies. Consequently specific grooming patterns emerge. In despotic species focusing grooming activity on preferred social partners significantly ameliorates GC levels in females of all ranks. In this study we investigated grooming patterns and GC management in Barbary macaques, a comparably relaxed species. We monitored changes in grooming behavior and cortisol (C) for females of different ranks. Our results show that the C‐amelioration associated with different grooming patterns had a gradual connection with dominance hierarchy: while higher‐ranking individuals showed lowest urinary C measures when they focused their grooming on selected partners within their social network, lower‐ranking individuals expressed lowest C levels when dispersing their grooming activity evenly across their social partners. We argue that the relatively relaxed social style of Barbary macaque societies allows individuals to flexibly adapt grooming patterns, which is associated with rank‐specific GC management. Am. J. Primatol. 77:688–700, 2015

Abstract

Aims/hypothesis Several forkhead box (FOX) transcription factor family members have important roles in controlling pancreatic cell fates and maintaining beta cell mass and function, including FOXA1, FOXA2 and FOXM1. In this study we have examined the importance of FOXP1, FOXP2 and FOXP4 of the FOXP subfamily in islet cell development and function. Methods Mice harbouring floxed alleles for Foxp1, Foxp2 and Foxp4 were crossed with pan-endocrine Pax6-Cre transgenic mice to generate single and compound Foxp mutant mice. Mice were monitored for changes in glucose tolerance by IPGTT, serum insulin and glucagon levels by radioimmunoassay, and endocrine cell development and proliferation by immunohistochemistry. Gene expression and glucose-stimulated hormone secretion experiments were performed with isolated islets. Results Only the triple-compound Foxp1/2/4 conditional knockout (cKO) mutant had an overt islet phenotype, manifested physiologically by hypoglycaemia and hypoglucagonaemia. This resulted from the reduction in glucagon-secreting alpha cell mass and function. The proliferation of alpha cells was profoundly reduced in Foxp1/2/4 cKO islets through the effects on mediators of replication (i.e. decreased Ccna2, Ccnb1 and Ccnd2 activators, and increased Cdkn1a inhibitor). Adult islet Foxp1/2/4 cKO beta cells secrete insulin normally while the remaining alpha cells have impaired glucagon secretion. Conclusions/interpretation Collectively, these findings reveal an important role for the FOXP1, 2, and 4 proteins in governing postnatal alpha cell expansion and function.

Abstract

E-Prime® is the leading software suite by Psychology Software Tools for designing and running Psychology lab experiments. The E-Primer is the perfect accompanying guide: It provides all the necessary knowledge to make E-Prime accessible to everyone. You can learn the tools of Psychological science by following the E-Primer through a series of entertaining, step-by-step recipes that recreate classic experiments. The updated E-Primer expands its proven combination of simple explanations, interesting tutorials and fun exercises, and makes even the novice student quickly confident to create their dream experiment.

Abstract

Odors are often difficult to identify and name, which leaves them vulnerable to the influence of language. The present study tests the boundaries of the effect of language on odor cognition by examining the effect of grammatical gender. We presented participants with male and female fragrances paired with descriptions of masculine or feminine grammatical gender. In Experiment 1 we found that memory for fragrances was enhanced when the grammatical gender of a fragrance description matched the gender of the fragrance. In Experiment 2 we found memory for fragrances was affected by both grammatical gender and gender associations in fragrance descriptions – recognition memory for odors was higher when the gender was incongruent. In sum, we demonstrated that even subtle aspects of language can affect odor cognition.

Abstract

Metaphor allows us to think and talk about one thing in terms of another, ratcheting up our cognitive and expressive capacity. It gives us concrete terms for abstract phenomena, for example, ideas become things we can grasp or let go of. Perceptual experience—characterised as physical and relatively concrete—should be an ideal source domain in metaphor, and a less likely target. But is this the case across diverse languages? And are some sensory modalities perhaps more concrete than others? This volume presents critical new data on perception metaphors from over 40 languages, including many which are under-studied. Aside from the wealth of data from diverse languages—modern and historical; spoken and signed—a variety of methods (e.g., natural language corpora, experimental) and theoretical approaches are brought together. This collection highlights how perception metaphor can offer both a bedrock of common experience and a source of continuing innovation in human communication

Abstract

The problem of identifying and correcting confusibles, i.e. context-sensitive spelling errors, in text is typically tackled using specifically trained machine learning classifiers. For each different set of confusibles, a specific classifier is trained and tuned. In this research, we investigate a more generic approach to context-sensitive confusible correction. Instead of using specific classifiers, we use one generic classifier based on a language model. This measures the likelihood of sentences with different possible solutions of a confusible in place. The advantage of this approach is that all confusible sets are handled by a single model. Preliminary results show that the performance of the generic classifier approach is only slightly worse that that of the specific classifier approach

Abstract

In the context of confusible disambiguation (spelling correction that requires context), the synchronous back-off strategy combined with traditional n-gram language models performs well. However, when alternatives consist of a different number of tokens, this classification technique cannot be applied directly, because the computation of the probabilities is skewed. Previous work already showed that probabilities based on different order n-grams should not be compared directly. In this article, we propose new probability metrics in which the size of the n is varied according to the number of tokens of the confusible alternative. This requires access to n-grams of variable length. Results show that the synchronous back-off method is extremely robust. We discuss the use of suffix trees as a technique to store variable length n-gram information efficiently.

Abstract

Two word-by-word, self-paced reading experiments investigated the speed with which readers were sensitive to discourse-level anomalies. An account arguing for delayed sensitivity (Guzman & Klin, 2000 Guzman, A. E. and Klin, C. M. 2000. Maintaining global coherence in reading: The role of sentence boundaries.. Memory & Cognition, 28: 722–730. [PubMed], [Web of Science ®], [Google Scholar]) was contrasted with one allowing for rapid sensitivity (Myers & O'Brien, 1998 Myers, J. L. and O'Brien, E. J. 1998. Accessing the discourse representation during reading.. Discourse Processes, 26: 131–157. [Taylor & Francis Online], [Web of Science ®], [Google Scholar]). Anomalies related to spatial information (Experiment 1) and character-attribute information (Experiment 2) were examined. Both experiments found that readers displayed rapid sensitivity to the anomalous information. A reading time penalty was observed for the region of text containing the anomalous information. This finding is most compatible with an account of text processing whereby incoming words are rapidly evaluated with respect to prior context. They are not consistent with an account that argues for delayed integration. Results are discussed in light of their implications for competing models of text processing.

Abstract

The ability to represent conditional information is central to human cognition. In two self-paced reading experiments we investigated how readers process counterfactual conditionals (e.g., If Darren had been athletic, he could probably have played on the rugby team ) and indicative conditionals (e.g., If Darren is athletic, he probably plays on the rugby team ). In Experiment 1 we focused on how readers process counterfactual conditional sentences. We found that processing of the antecedent of counterfactual conditionals was rapidly constrained by prior context (i.e., knowing whether Darren was or was not athletic). A reading-time penalty was observed for the critical region of text comprising the last word of the antecedent and the first word of the consequent when the information in the antecedent did not fit with prior context. In Experiment 2 we contrasted counterfactual conditionals with indicative conditionals. For counterfactual conditionals we found the same effect on the critical region as we found in Experiment 1. In contrast, however, we found no evidence that processing of the antecedent of indicative conditionals was constrained by prior context. For indicative conditionals (but not for counterfactual conditionals), the results we report are consistent with the suppositional account of conditionals. We propose that current theories of conditionals need to be able to account for online processing differences between indicative and counterfactual conditionals

Abstract

Informal verbal interaction is the core matrix for human social life. A mechanism for coordinating this basic mode of interaction is a system of turn-taking that regulates who is to speak and when. Yet relatively little is known about how this system varies across cultures. The anthropological literature reports significant cultural differences in the timing of turn-taking in ordinary conversation. We test these claims and show that in fact there are striking universals in the underlying pattern of response latency in conversation. Using a worldwide sample of 10 languages drawn from traditional indigenous communities to major world languages, we show that all of the languages tested provide clear evidence for a general avoidance of overlapping talk and a minimization of silence between conversational turns. In addition, all of the languages show the same factors explaining within-language variation in speed of response. We do, however, find differences across the languages in the average gap between turns, within a range of 250 ms from the cross-language mean. We believe that a natural sensitivity to these tempo differences leads to a subjective perception of dramatic or even fundamental differences as offered in ethnographic reports of conversational style. Our empirical evidence suggests robust human universals in this domain, where local variations are quantitative only, pointing to a single shared infrastructure for language use with likely ethological foundations.

Abstract

Bilingual children are often exposed to non-native speech through their parents. Yet, little is known about the relation between bilingual preschoolers’ speech production and their speech input. The present study investigated the production of voice onset time (VOT) by Dutch-German bilingual preschoolers and their sequential bilingual mothers. The findings reveal an association between maternal VOT and bilingual children’s VOT in the heritage language German as well as in the majority language Dutch. By contrast, no input-production association was observed in the VOT production of monolingual German-speaking children and monolingual Dutch-speaking children. The results of this study provide the first empirical evidence that non-native and attrited maternal speech contributes to the often-observed linguistic differences between bilingual children and their monolingual peers.

Abstract

Finnish vowel harmony rules require that if the vowel in the first syllable of a word belongs to one of two vowel sets, then all subsequent vowels in that word must belong either to the same set or to a neutral set. A harmony mismatch between two syllables containing vowels from the opposing sets thus signals a likely word boundary. We report five experiments showing that Finnish listeners can exploit this information in an on-line speech segmentation task. Listeners found it easier to detect words likehymyat the end of the nonsense stringpuhymy(where there is a harmony mismatch between the first two syllables) than in the stringpyhymy(where there is no mismatch). There was no such effect, however, when the target words appeared at the beginning of the nonsense string (e.g.,hymypuvshymypy). Stronger harmony effects were found for targets containing front harmony vowels (e.g.,hymy) than for targets containing back harmony vowels (e.g.,paloinkypaloandkupalo). The same pattern of results appeared whether target position within the string was predictable or unpredictable. Harmony mismatch thus appears to provide a useful segmentation cue for the detection of word onsets in Finnish speech.

Abstract

In this study the N400 component of the event-related potential was used to investigate spoken sentence understanding in Broca's and Wernicke's aphasics. The aim of the study was to determine whether spoken sentence comprehension problems in these patients might result from a deficit in the on-line integration of lexical information. Subjects listened to sentences spoken at a normal rate. In half of these sentences, the meaning of the final word of the sentence matched the semantic specifications of the preceding sentence context. In the other half of the sentences, the sentence-final word was anomalous with respect to the preceding sentence context. The N400 was measured to the sentence-final words in both conditions. The results for the aphasic patients (n = 14) were analyzed according to the severity of their comprehension deficit and compared to a group of 12 neurologically unimpaired age-matched controls, as well as a group of 6 nonaphasic patients with a lesion in the right hemisphere. The nonaphasic brain damaged patients and the aphasic patients with a light comprehension deficit (high comprehenders, n = 7) showed an N400 effect that was comparable to that of the neurologically unimpaired subjects. In the aphasic patients with a moderate to severe comprehension deficit (low comprehenders, n = 7), a reduction and delay of the N400 effect was obtained. In addition, the P300 component was measured in a classical oddball paradigm, in which subjects were asked to count infrequent low tones in a random series of high and low tones. No correlation was found between the occurrence of N400 and P300 effects, indicating that changes in the N400 results were related to the patients' language deficit. Overall, the pattern of results was compatible with the idea that aphasic patients with moderate to severe comprehension problems are impaired in the integration of lexical information into a higher order representation of the preceding sentence context.

Abstract

There exists a clear association between animacy and the grammatical function of transitive subject. The grammar of some languages require the transitive subject to be high in animacy, or at least higher than the object. A similar animacy preference has been observed in processing studies in languages without such a categorical animacy effect. This animacy preference has been mainly established in structures in which either one or both arguments are provided before the verb. Our goal was to establish (i) whether this preference can already be observed before any argument is provided, and (ii) whether this preference is mediated by verbal information. To this end we exploited the V2 property of Dutch which allows the verb to precede its arguments. Using a visual-world eye-tracking paradigm we presented participants with V2 structures with either an auxiliary (e.g. Gisteren heeft X … ‘Yesterday, X has …’) or a lexical main verb (e.g. Gisteren motiveerde X … ‘Yesterday, X motivated …’) and we measured looks to the animate referent. The results indicate that the animacy preference can already be observed before arguments are presented and that the selectional restrictions of the verb mediate this bias, but do not override it completely.

Abstract

General knowledge acquisition entails the extraction of statistical regularities from the environment. At high levels of complexity, this may involve the extraction, and consolidation, of associative regularities across event memories. The underlying neural mechanisms would likely involve a hippocampo-neocortical dialog, as proposed previously for system-level consolidation. To test these hypotheses, we assessed possible differences in consolidation between associative memories containing cross-episodic regularities and unique associative memories. Subjects learned face–location associations, half of which responded to complex regularities regarding the combination of facial features and locations, whereas the other half did not. Importantly, regularities could only be extracted over hippocampus-encoded, associative aspects of the items. Memory was assessed both immediately after encoding and 48 h later, under fMRI acquisition. Our results suggest that processes related to system-level reorganization occur preferentially for regular associations across episodes. Moreover, the build-up of general knowledge regarding regular associations appears to involve the coordinated activity of the hippocampus and mediofrontal regions. The putative cross-talk between these two regions might support a mechanism for regularity extraction. These findings suggest that the consolidation of cross-episodic regularities may be a key mechanism underlying general knowledge acquisition.

Abstract

Five word-spotting experiments explored the role of consonantal and vocalic phonotactic cues in the segmentation of spoken Italian. The first set of experiments tested listeners' sensitivity to phonotactic constraints cueing syllable boundaries. Participants were slower in spotting words in nonsense strings when target onsets were misaligned (e.g., lago in ri.blago) than when they were aligned (e.g., lago in rin.lago) with phonotactically determined syllabic boundaries. This effect held also for sequences that occur only word-medially (e.g., /tl/ in ri.tlago), and competition effects could not account for the disadvantage in the misaligned condition. Similarly, target detections were slower when their offsets were misaligned (e.g., cittaacute in cittaacuteu.ba) than when they were aligned (e.g., cittaacute in cittaacute.oba) with a phonotactic syllabic boundary. The second set of experiments tested listeners' sensitivity to phonotactic cues, which specifically signal lexical (and not just syllable) boundaries. Results corroborate the role of syllabic information in speech segmentation and suggest that Italian listeners make little use of additional phonotactic information that specifically cues word boundaries.

Abstract

Two cross-modal priming experiments tested whether lexical access is constrained by syllabic structure in Italian. Results extend the available Italian data on the processing of stressed syllables showing that syllabic information restricts the set of candidates to those structurally consistent with the intended word (Experiment 1). Lexical access, however, takes place as soon as possible and it is not delayed till the incoming input corresponds to the first syllable of the word. And, the initial activated set includes candidates whose syllabic structure does not match the intended word (Experiment 2). The present data challenge the early hypothesis that in Romance languages syllables are the units for lexical access during spoken word recognition. The implications of the results for our understanding of the role of syllabic information in language processing are discussed.

Abstract

Memory representations of words are thought to undergo changes with consolidation: Episodic memories of novel words are transformed into lexical representations that interact with other words in the mental dictionary. Behavioral studies have shown that this lexical integration process is enhanced when there is more time for consolidation. Neuroimaging studies have further revealed that novel word representations are initially represented in a hippocampally-centered system, whereas left posterior middle temporal cortex activation increases with lexicalization. In this study, we measured behavioral and brain responses to newly-learned words in children. Two groups of Dutch children, aged between 8-10 and 14-16 years, were trained on 30 novel Japanese words depicting novel concepts. Children were tested on word-forms, word-meanings, and the novel words’ influence on existing word processing immediately after training, and again after a week. In line with the adult findings, hippocampal involvement decreased with time. Lexical integration, however, was not observed immediately or after a week, neither behaviorally nor neurally. It appears that time alone is not always sufficient for lexical integration to occur. We suggest that other factors (e.g., the novelty of the concepts and familiarity with the language the words are derived from) might also influence the integration process.

Abstract

The aim of the present study was to examine whether repetition of radicals during training of Chinese characters leads to better word acquisition performance in beginning learners of Chinese as a foreign language. Thirty Dutch university students were trained on 36 Chinese one-character words for their pronunciations and meanings. They were also exposed to the specifics of the radicals, that is, for phonetic radicals, the associated pronunciation was explained, and for semantic radicals the associated categorical meanings were explained. Results showed that repeated exposure to phonetic and semantic radicals through character pronunciation and meaning trainings indeed induced better understanding of those radicals that were shared among different characters. Furthermore, characters in the training set that shared phonetic radicals were pronounced better than those that did not. Repetition of semantic radicals across different characters, however, hindered the learning of exact meanings. Students generally confused the meanings of other characters that shared the semantic radical. The study shows that in the initial stage of learning, overlapping information of the shared radicals are effectively learned. Acquisition of the specifics of individual characters, however, requires more training.

Abstract

The rates of overspecification of color, pattern, and size are compared, to investigate how salience and absoluteness contribute to the production of overspecification. Color and pattern are absolute and salient attributes, whereas size is relative and less salient. Additionally, a tendency toward consistent responses is assessed. Using a within-participants design, we find similar rates of color and pattern overspecification, which are both higher than the rate of size overspecification. Using a between-participants design, however, we find similar rates of pattern and size overspecification, which are both lower than the rate of color overspecification. This indicates that although many speakers are more likely to include color than pattern (probably because color is more salient), they may also treat pattern like color due to a tendency toward consistency. We find no increase in size overspecification when the salience of size is increased, suggesting that speakers are more likely to include absolute than relative attributes. However, we do find an increase in size overspecification when mentioning the attributes is triggered, which again shows that speakers tend to refer in a consistent manner, and that there are circumstances in which even size overspecification is frequently produced.

Abstract

Although visual imagery is argued to be an essential component of autobiographical memory, there have been surprisingly few studies on autobiographical memory processes in blind individuals, who have had no or limited visual input. The purpose of the present study was to investigate how blindness affects retrieval and phenomenology of autobiographical memories. We asked 48 congenital/early blind and 48 sighted participants to recall autobiographical memories in response to six cue words, and to fill out the Autobiographical Memory Questionnaire measuring a number of variables including imagery, belief and recollective experience associated with each memory. Blind participants retrieved fewer memories and reported higher auditory imagery at retrieval than sighted participants. Moreover, within the blind group, participants with total blindness reported higher auditory imagery than those with some light perception. Blind participants also assigned higher importance, belief and recollection ratings to their memories than sighted participants. Importantly, these group differences remained the same for recent as well as childhood memories.

Abstract

The neural processing of speech leads to specific patterns in the brain which can be measured as, e.g., EEG signals. When properly aligned with the speech input and averaged over many tokens, the Event Related Potential (ERP) signal is able to differentiate specific contrasts between speech signals. Well-known effects relate to the difference between expected and unexpected words, in particular in the N400, while effects in N100 and P200 are related to attention and acoustic onset effects. Most EEG studies deal with the amplitude of EEG signals over time, sidestepping the effect of phase and phase synchronization. This paper investigates the relation between phase in the EEG signals measured in an auditory lexical decision task by Dutch participants listening to full and reduced English word forms. We show that phase synchronization takes place across stimulus conditions, and that the so-called circular variance is narrowly related to the type of contrast between stimuli.

Abstract

This paper presents DIANA, a new computational model of human speech processing. It is the first model that simulates the complete processing chain from the on-line processing of an acoustic signal to the execution of a response, including reaction times. Moreover it assumes minimal modularity. DIANA consists of three components. The activation component computes a probabilistic match between the input acoustic signal and representations in DIANA’s lexicon, resulting in a list of word hypotheses changing over time as the input unfolds. The decision component operates on this list and selects a word as soon as sufficient evidence is available. Finally, the execution component accounts for the time to execute a behavioral action. We show that DIANA well simulates the average participant in a word recognition experiment.

Abstract

DIANA is an end-to-end computational model of speech processing, which takes as input the speech signal, and provides as output the orthographic transcription of the stimulus, a word/non-word judgment and the associated estimated reaction time. So far, the model has only been tested for Dutch. In this paper, we extend DIANA such that it can also process North American English. The model is tested by having it simulate human participants in a large scale North American English lexical decision experiment. The simulations show that DIANA can adequately approximate the reaction times of an average participant (r = 0.45). In addition, they indicate that DIANA does not yet adequately model the cognitive processes that take place after stimulus offset.

Abstract

Temporal cues can be used to selectively attend to relevant information during abundant sensory stimulation. However, such cues differ vastly in the accuracy of their temporal estimates, ranging from very predictable to very unpredictable. When cues are strongly predictable, attention may facilitate selective processing by aligning relevant incoming information to high neuronal excitability phases of ongoing low-frequency oscillations. However, top-down effects on ongoing oscillations when temporal cues have some predictability, but also contain temporal uncertainties, are unknown. Here, we experimentally created such a situation of mixed predictability and uncertainty: A target could occur within a limited time window after cue but was always unpredictable in exact timing. Crucially to assess top-down effects in such a mixed situation, we manipulated target probability. High target likelihood, compared with low likelihood, enhanced delta oscillations more strongly as measured by evoked power and intertrial coherence. Moreover, delta phase modulated detection rates for probable targets. The delta frequency range corresponds with half-a-period to the target occurrence window and therefore suggests that low-frequency phase reset is engaged to produce a long window of high excitability when event timing is uncertain within a restricted temporal window.

Abstract

The brain is inherently proactive, constantly predicting the when (moment) and what (content) of future input in order to optimize information processing. Previous research on such predictions has mainly studied the “when” or “what” domain separately, missing to investigate the potential integration of both types of predictive information. In the absence of such integration, temporal cues are assumed to enhance any upcoming content at the predicted moment in time (general temporal predictor). However, if the when and what prediction domain were integrated, a much more flexible neural mechanism may be proposed in which temporal-feature interactions would allow for the creation of multiple concurrent time-content predictions (parallel time-content predictor). Here, we used a temporal association paradigm in two experiments in which sound identity was systematically paired with a specific time delay after the offset of a rhythmic visual input stream. In Experiment 1, we revealed that participants associated the time delay of presentation with the identity of the sound. In Experiment 2, we unexpectedly found that the strength of this temporal association was negatively related to the EEG steady-state evoked responses (SSVEP) in preceding trials, showing that after high neuronal responses participants responded inconsistent with the time-content associations, similar to adaptation mechanisms. In this experiment, time-content associations were only present for low SSVEP responses in previous trials. These results tentatively show that it is possible to represent multiple time-content paired predictions in parallel, however, future research is needed to investigate this interaction further.

Abstract

The role of oscillatory phase for perceptual and cognitive processes is being increasingly acknowledged. To date, little is known about the direct role of phase in categorical perception. Here we show in two separate experiments that the identification of ambiguous syllables that can either be perceived as / da/ or / ga/ is biased by the underlying oscillatory phase as measured with EEG and sensory entrainment to rhythmic stimuli. The measured phase difference in which perception is biased toward / da/ or / ga/ exactly matched the different temporal onset delays in natural audiovisual speech between mouth movements and speech sounds, which last 80 ms longer for / ga/ than for / da/. These results indicate the functional relationship between prestimulus phase and syllable identification, and signify that the origin of this phase relationship could lie in exposure and subsequent learning of unique audiovisual temporal onset differences.

Abstract

In everyday life, people see, describe and remember motion events. We tested whether the type of motion event information (path or manner) encoded in speech and gesture predicts which information is remembered and if this varies across speakers of typologically different languages. We focus on intransitive motion events (e.g., a woman running to a tree) that are described differently in speech and co-speech gesture across languages, based on how these languages typologically encode manner and path information (Kita & Özyürek, 2003; Talmy, 1985). Speakers of Dutch (n = 19) and Turkish (n = 22) watched and described motion events. With a surprise (i.e. unexpected) recognition memory task, memory for manner and path components of these events was measured. Neither Dutch nor Turkish speakers’ memory for manner went above chance levels. However, we found a positive relation between path speech and path change detection: participants who described the path during encoding were more accurate at detecting changes to the path of an event during the memory task. In addition, the relation between path speech and path memory changed with native language: for Dutch speakers encoding path in speech was related to improved path memory, but for Turkish speakers no such relation existed. For both languages, co-speech gesture did not predict memory speakers. We discuss the implications of these findings for our understanding of the relations between speech, gesture, type of encoding in language and memory.

Abstract

Apraxia of Speech (AOS) is a motor speech disorder whose precise nature is still poorly understood. A recent behavioural experiment featuring a noise masking paradigm suggests that AOS reflects a disruption of feedforward control, whereas feedback control is spared and plays a more prominent role in achieving and maintaining segmental contrasts [10]. In the present study, we set out to validate the interpretation of AOS as a feedforward impairment by means of a series of computational simulations with the DIVA model [6, 7] mimicking the behavioural experiment. Simulation results showed a larger reduction in vowel spacing and a smaller vowel dispersion in the masking condition compared to the no-masking condition for the simulated feedforward deficit, whereas the other groups showed an opposite pattern. These results mimic the patterns observed in the human data, corroborating the notion that AOS can be conceptualized as a deficit in feedforward control

Abstract

Difficulties with pragmatic aspects of communication are universal across individuals with autism spectrum disorders (ASDs). Here we focused on an aspect of pragmatic language comprehension that is relevant to social interaction in daily life: the integration of speaker characteristics inferred from the voice with the content of a message. Using functional magnetic resonance imaging (fMRI), we examined the neural correlates of the integration of voice-based inferences about the speaker’s age, gender or social background, and sentence content in adults with ASD and matched control participants. Relative to the control group, the ASD group showed increased activation in right inferior frontal gyrus (RIFG; Brodmann area 47) for speakerincongruent sentences compared to speaker-congruent sentences. Given that both groups performed behaviourally at a similar level on a debriefing interview outside the scanner, the increased activation in RIFG for the ASD group was interpreted as being compensatory in nature. It presumably reflects spill-over processing from the language dominant left hemisphere due to higher task demands faced by the participants with ASD when integrating speaker characteristics and the content of a spoken sentence. Furthermore, only the control group showed decreased activation for speaker-incongruent relative to speaker-congruent sentences in right ventral medial prefrontal cortex (vMPFC; Brodmann area 10), including right anterior cingulate cortex (ACC; Brodmann area 24/32). Since vMPFC is involved in self-referential processing related to judgments and inferences about self and others, the absence of such a modulation in vMPFC activation in the ASD group possibly points to atypical default self-referential mental activity in ASD. Our results show that in ASD compensatory mechanisms are necessary in implicit, low-level inferential processes in spoken language understanding. This indicates that pragmatic language problems in ASD are not restricted to high-level inferential processes, but encompass the most basic aspects of pragmatic language processing.

Abstract

When interpreting a message, a listener takes into account several sources of linguistic and extralinguistic information. Here we focused on one particular form of extralinguistic information, certain speaker characteristics as conveyed by the voice. Using functional magnetic resonance imaging, we examined the neural structures involved in the unification of sentence meaning and voice-based inferences about the speaker's age, sex, or social background. We found enhanced activation in the inferior frontal gyrus bilaterally (BA 45/47) during listening to sentences whose meaning was incongruent with inferred speaker characteristics. Furthermore, our results showed an overlap in brain regions involved in unification of speaker-related information and those used for the unification of semantic and world knowledge information [inferior frontal gyrus bilaterally (BA 45/47) and left middle temporal gyrus (BA 21)]. These findings provide evidence for a shared neural unification system for linguistic and extralinguistic sources of information and extend the existing knowledge about the role of inferior frontal cortex as a crucial component for unification during language comprehension.

Abstract

Purpose: The question of how and when English-speaking children acquire auxiliaries is the subject of extensive debate. Some researchers posit the existence of innately given Universal Grammar principles to guide acquisition, although some aspects of the auxiliary system must be learned from the input. Others suggest that auxiliaries can be learned without Universal Grammar, citing evidence of piecemeal learning in their support. This study represents a unique attempt to trace the development of auxiliary syntax by using a longitudinal elicitation methodology. Method: Twelve English-speaking children participated in 3 tasks designed to elicit auxiliary BE in declaratives and yes/no and wh-questions. They completed each task 6 times in total between the ages of 2;10 (years;months) and 3;6. Results: The children’s levels of correct use of 2 forms of BE (is,are) differed according to auxiliary form and sentence structure, and these relations changed over development. An analysis of the children’s errors also revealed complex interactions between these factors. Conclusion: These data are problematic for existing accounts of auxiliary acquisition and highlight the need for researchers working within both generativist and constructivist frameworks to develop more detailed theories of acquisition that directly predict the pattern of acquisition observed.

Abstract

Our understanding of the functioning of the brain is primarily based on an average model of the brain's functional organisation, and any deviation from the standard is considered as random noise or a pathological appearance. Studying pathologies has, however, greatly contributed to our understanding of brain functions. For instance, the study of naturally-occurring or surgically-induced brain lesions revealed that language is predominantly lateralised to the left hemisphere while perception/action and emotion are commonly lateralised to the right hemisphere. The lateralisation of function was subsequently replicated by task-related functional neuroimaging in the healthy population. Despite its high significance and reproducibility, this pattern of lateralisation of function is true for most, but not all participants. Bilateral and flipped representations of classically lateralised functions have been reported during development and in the healthy adult population for language, perception/action and emotion. Understanding these different functional representations at an individual level is crucial to improve the sophistication of our models and account for the variance in developmental trajectories, cognitive performance differences and clinical recovery. With the availability of in vivo neuroimaging, it has become feasible to study large numbers of participants and reliably characterise individual differences, also referred to as phenotypes. Yet, we are at the beginning of inter-individual variability modelling, and new theories of brain function will have to account for these differences across participants.

Abstract

To test the hypothesis that thalamic midline nuclei play a transient role in memory consolidation, we reanalyzed a prospective functional MRI study, contrasting recent and progressively more remote memory retrieval. We revealed a transient thalamic connectivity increase with the hippocampus, the medial prefrontal cortex (mPFC), and a parahippocampal area, which decreased with time. In turn, mPFC-parahippocampal connectivity increased progressively. These findings support a model in which thalamic midline nuclei serve as a hub linking hippocampus, mPFC, and posterior representational areas during memory retrieval at an early (2 h) stage of consolidation, extending classical systems consolidation models by attributing a transient role to midline thalamic nuclei.

Abstract

Infants expect people to direct actions toward objects, and they respond to actions directed to themselves, but do they have expectations about actions directed to third parties? In two experiments, we used eye tracking to investigate 1- and 2-year-olds’ expectations about communicative actions addressed to a third party. Experiment 1 presented infants with videos where an adult (the Emitter) either uttered a sentence or produced non-speech sounds. The Emitter was either face-to-face with another adult (the Recipient) or the two were back-to-back. The Recipient did not respond to any of the sounds. We found that 2-, but not 1-year-olds looked quicker and longer at the Recipient following speech than non-speech, suggesting that they expected her to respond to speech. These effects were specific to the face-to-face context. Experiment 2 presented 1-year-olds with similar face-to-face exchanges but modified to engage infants and minimize task demands. The infants looked quicker to the Recipient following speech than non-speech, suggesting that they expected a response to speech. The study suggests that by 1 year of age infants expect communicative actions to be directed at a third-party listener.

Abstract

Germline mutations in PTEN, which encodes a widely expressed phosphatase, was mapped to 10q23 and identified as the susceptibility gene for Cowden syndrome, characterized by macrocephaly and high risks of breast, thyroid, and other cancers. The phenotypic spectrum of PTEN mutations expanded to include autism with macrocephaly only 10 years ago. Neurological studies of patients with PTEN-associated autism spectrum disorder (ASD) show increases in cortical white matter and a distinctive cognitive profile, including delayed language development with poor working memory and processing speed. Once a germline PTEN mutation is found, and a diagnosis of phosphatase and tensin homolog (PTEN) hamartoma tumor syndrome made, the clinical outlook broadens to include higher lifetime risks for multiple cancers, beginning in childhood with thyroid cancer. First described as a tumor suppressor, PTEN is a major negative regulator of the phosphatidylinositol 3-kinase/protein kinase B/mammalian target of rapamycin (mTOR) signaling pathway—controlling growth, protein synthesis, and proliferation. This canonical function combines with less well-understood mechanisms to influence synaptic plasticity and neuronal cytoarchitecture. Several excellent mouse models of Pten loss or dysfunction link these neural functions to autism-like behavioral abnormalities, such as altered sociability, repetitive behaviors, and phenotypes like anxiety that are often associated with ASD in humans. These models also show the promise of mTOR inhibitors as therapeutic agents capable of reversing phenotypes ranging from overgrowth to low social behavior. Based on these findings, therapeutic options for patients with PTEN hamartoma tumor syndrome and ASD are coming into view, even as new discoveries in PTEN biology add complexity to our understanding of this master regulator.

Abstract

Synaesthesia is a neurological phenomenon affecting perception, where triggering stimuli (e.g. letters and numbers) elicit unusual secondary sensory experiences (e.g. colours). Family-based studies point to a role for genetic factors in the development of this trait. However, the contributions of common genomic variation to synaesthesia have not yet been investigated. Here, we present the SynGenes cohort, the largest genotyped collection of unrelated people with grapheme–colour synaesthesia (n = 723). Synaesthesia has been associated with a range of other neuropsychological traits, including enhanced memory and mental imagery, as well as greater sensory sensitivity. Motivated by the prior literature on putative trait overlaps, we investigated polygenic scores derived from published genome-wide scans of schizophrenia and autism spectrum disorder (ASD), comparing our SynGenes cohort to 2181 non-synaesthetic controls. We found a very slight association between schizophrenia polygenic scores and synaesthesia (Nagelkerke's R2 = 0.0047, empirical p = 0.0027) and no significant association for scores related to ASD (Nagelkerke's R2 = 0.00092, empirical p = 0.54) or body mass index (R2 = 0.00058, empirical p = 0.60), included as a negative control. As sample sizes for studying common genomic variation continue to increase, genetic investigations of the kind reported here may yield novel insights into the shared biology between synaesthesia and other traits, to complement findings from neuropsychology and brain imaging.

Abstract

Peak bone mass achieved in adolescence is a determinant of bone mass in later life. In order to identify genetic variants affecting bone mineral density (BMD), we performed a genome-wide association study of BMD and related traits in 1518 children from the Avon Longitudinal Study of Parents and Children (ALSPAC). We compared results with a scan of 134 adults with high or low hip BMD. We identified associations with BMD in an area of chromosome 12 containing the Osterix (SP7) locus, a transcription factor responsible for regulating osteoblast differentiation (ALSPAC: P = 5.8 x 10(-4); Australia: P = 3.7 x 10(-4)). This region has previously shown evidence of association with adult hip and lumbar spine BMD in an Icelandic population, as well as nominal association in a UK population. A meta-analysis of these existing studies revealed strong association between SNPs in the Osterix region and adult lumbar spine BMD (P = 9.9 x 10(-11)). In light of these findings, we genotyped a further 3692 individuals from ALSPAC who had whole body BMD and confirmed the association in children as well (P = 5.4 x 10(-5)). Moreover, all SNPs were related to height in ALSPAC children, but not weight or body mass index, and when height was included as a covariate in the regression equation, the association with total body BMD was attenuated. We conclude that genetic variants in the region of Osterix are associated with BMD in children and adults probably through primary effects on growth.

Abstract

The neural response to a stimulus is influenced by endogenous factors such as expectation and attention. Current research suggests that expectation and attention exert their effects in opposite directions, where expectation decreases neural activity in sensory areas, while attention increases it. However, expectation and attention are usually studied either in isolation or confounded with each other. A recent study suggests that expectation and attention may act jointly on sensory processing, by increasing the neural response to expected events when they are attended, but decreasing it when they are unattended. Here we test this hypothesis in an auditory temporal cueing paradigm using magnetoencephalography in humans. In our study participants attended to, or away from, tones that could arrive at expected or unexpected moments. We found a decrease in auditory beta band synchrony to expected (versus unexpected) tones if they were unattended, but no difference if they were attended. Modulations in beta power were already evident prior to the expected onset times of the tones. These findings suggest that expectation and attention jointly modulate sensory processing.

Abstract

In this study, we investigate the timing of pre-answer inbreaths in order to shed light on the time course of response planning and execution in conversational turn-taking. Using acoustic and inductive plethysmography recordings of seven dyadic conversations in Dutch, we show that pre-answer inbreaths in conversation typically begin briefly after the end of questions. We also show that the presence of a pre-answer inbreath usually co-occurs with substantially delayed answers, with a modal latency of 576 ms vs. 100 ms for answers not preceded by an inbreath. Based on previously reported minimal latencies for internal intercostal activation and the production of speech sounds, we propose that vocal responses, either in the form of a pre-utterance inbreath or of speech proper when an inbreath is not produced, are typically launched in reaction to information present in the last portion of the interlocutor’s turn. We also show that short responses are usually made on residual breath, while longer responses are more often preceded by an inbreath. This relation of inbreaths to answer length suggests that by the time an inbreath is launched, typically during the last few hundred milliseconds of the question, the length of the answer is often prepared to some extent. Together, our findings are consistent with a two-stage model of response planning in conversational turn-taking: early planning of content often carried out in overlap with the incoming turn, and late launching of articulation based on the identification of turn-final cues

Abstract

This article describes how the tonal elements of two common Spanish intonation contours –the falling statement and the low-rising-falling request– align with the segmental string in broad-focus utterances differing in number of prosodic words. Using an imitation-and-completion task, we show that (i) the last stressed syllable of the utterance, traditionally viewed as carrying the ‘nuclear’ accent, associates with either a high or a low tonal element depending on phrase length (ii) that certain tonal elements can be realized or omitted depending on the availability of specific metrical positions in their intonational phrase, and (iii) that the high tonal element of the request contour associates with either a stressed syllable or an intonational phrase edge depending on phrase length. On the basis of these facts, and in contrast to previous descriptions of Spanish intonation relying on obligatory and constant nuclear contours (e.g., L* L% for all neutral statements), we argue for a less constrained intonational morphology involving tonal units linked to the segmental string via contour-specific principles.

Abstract

We investigate the extent to which French polar questions and continuation statements, two types of utterances with similar morphosyntactic and intonational forms but different pragmatic functions, can be distinguished in conversational data based on phonetic and visual bodily information. We show that the two utterance types can be distinguished well over chance level by automatic classification models including several phonetic and visual cues. We also show that a considerable amount of relevant phonetic and visual information is present before the last portion of the utterances, potentially assisting early speech act recognition by addressees. These findings indicate that bottom-up phonetic and visual cues may play an important role during the production and recognition of speech acts alongside top-down contextual information.

Abstract

The present study shows that [t] consonants are affected by probabilistic factors in a syllable-timed language as French, and in spontaneous as well as in journalistic speech. Study 1 showed a word bigram frequency effect in spontaneous French, but its exact nature depended on the corpus on which the probabilistic measures were based. Study 2 investigated journalistic speech and showed an effect of the joint frequency of the test word and its following word. We discuss the possibility that these probabilistic effects are due to the speaker’s planning of upcoming words, and to the speaker’s adaptation to the listener’s needs.

Abstract

Violations of the maxims of Quantity occur when utterances provide more (over-specified) or less (under-specified) information than strictly required for referent identification. While behavioural datasuggest that under-specified expressions lead to comprehension difficulty and communicative failure, there is no consensus as to whether over-specified expressions are also detrimental to comprehension. In this study we shed light on this debate, providing neurophysiological evidence supporting the view that extra information facilitates comprehension. We further present novel evidence that referential failure due to under-specification is qualitatively different from explicit cases of referential failure, when no matching referential candidate is available in the context.

Abstract

Contrary to the Gricean maxims of quantity (Grice, in: Cole, Morgan (eds) Syntax and semantics: speech acts, vol III, pp 41–58, Academic Press, New York, 1975), it has been repeatedly shown that speakers often include redundant information in their utterances (over-specifications). Previous research on referential communication has long debated whether this redundancy is the result of speaker-internal or addressee-oriented processes, while it is also unclear whether referential redundancy hinders or facilitates comprehension. We present an information-theoretic explanation for the use of over-specification in visually-situated communication, which quantifies the amount of uncertainty regarding the referent as entropy (Shannon in Bell Syst Tech J 5:10, https://doi.org/10.1002/j.1538-7305.1948.tb01338.x, 1948). Examining both the comprehension and production of over-specifications, we present evidence that (a) listeners’ processing is facilitated by the use of redundancy as well as by a greater reduction of uncertainty early on in the utterance, and (b) that at least for some speakers, listeners’ processing concerns influence their encoding of over-specifications: Speakers were more likely to use redundant adjectives when these adjectives reduced entropy to a higher degree than adjectives necessary for target identification.

Abstract

Purpose: To determine clinical phenotypes, evolution and genetic background of a large family with a combination of two unusual forms of reflex epilepsies. Method: Phenotyping was performed in eighteen family members (10 F, 8 M) including standardized EEG recordings with intermittent photic stimulation (IPS). Genetic analyses (linkage scans, Whole Exome Sequencing (WES) and Functional studies) were performed using photoparoxysmal EEG responses (PPRs) as affection status. Results: The proband suffered from speaking induced jaw-jerks and increasing limb jerks evoked by flickering sunlight since about 50 years of age. Three of her family members had the same phenotype. Generalized PPRs were found in seven members (six above 50 years of age) with myoclonus during the PPR. Evolution was typical: Sensitivity to lights with migraine-like complaints around adolescence, followed by jerks evoked by lights and spontaneously with dropping of objects, and strong increase of light sensitivity and onset of talking induced jaw jerks around 50 years. Linkage analysis showed suggestive evidence for linkage to four genomic regions. All photosensitive family members shared a heterozygous R129C mutation in the SCNM1 gene that regulates splicing of voltage gated ion channels. Mutation screening of 134 unrelated PPR patients and 95 healthy controls, did not replicate these findings. Conclusion: This family presents a combination of two rare reflex epilepsies. Genetic analysis favors four genomic regions and points to a shared SCNM1 mutation that was not replicated in a general cohort of photosensitive subjects. Further genetic studies in families with similar combination of features are warranted. (C) 2015 British Epilepsy Association. Published by Elsevier Ltd. All rights reserved.

Abstract

Objective: Patients with Parkinson's disease (PD) often suffer from impairments in executive functions, such as working memory deficits. It is widely held that dopamine depletion in the striatum contributes to these impairments through decreased activity and connectivity between task-related brain networks. We investigated this hypothesis by studying task-related network activity and connectivity within a sample of de novo patients with PD, versus healthy controls, during a visuospatial working memory task. Methods: Sixteen de novo PD patients and 35 matched healthy controls performed a visuospatial n-back task while we measured their behavioral performance and neural activity using functional magnetic resonance imaging. We constructed regions-of-interest in the bilateral inferior parietal cortex (IPC), bilateral dorsolateral prefrontal cortex (DLPFC), and bilateral caudate nucleus to investigate group differences in task-related activity. We studied network connectivity by assessing the functional connectivity of the bilateral DLPFC and by assessing effective connectivity within the frontoparietal and the frontostriatal networks. Results: PD patients, compared with controls, showed trend-significantly decreased task accuracy, significantly increased task-related activity in the left DLPFC and a trend-significant increase in activity of the right DLPFC, left caudate nucleus, and left IPC. Furthermore, we found reduced functional connectivity of the DLPFC with other task-related regions, such as the inferior and superior frontal gyri, in the PD group, and group differences in effective connectivity within the frontoparietal network. Interpretation: These findings suggest that the increase in working memory-related brain activity in PD patients is compensatory to maintain behavioral performance in the presence of network deficits. Hum Brain Mapp 36:1554-1566, 2015. (c) 2015 Wiley Periodicals, Inc.

Abstract

Action, gesture, and sign represent unique aspects of human communication that use form and movement to convey meaning. Researchers typically use manual coding of video data to characterize naturalistic, meaningful movements at various levels of description, but the availability of markerless motion-tracking technology allows for quantification of the kinematic features of gestures or any meaningful human movement. We present a novel protocol for extracting a set of kinematic features from movements recorded with Microsoft Kinect. Our protocol captures spatial and temporal features, such as height, velocity, submovements/strokes, and holds. This approach is based on studies of communicative actions and gestures and attempts to capture features that are consistently implicated as important kinematic aspects of communication. We provide open-source code for the protocol, a description of how the features are calculated, a validation of these features as quantified by our protocol versus manual coders, and a discussion of how the protocol can be applied. The protocol effectively quantifies kinematic features that are important in the production (e.g., characterizing different contexts) as well as the comprehension (e.g., used by addressees to understand intent and semantics) of manual acts. The protocol can also be integrated with qualitative analysis, allowing fast and objective demarcation of movement units, providing accurate coding even of complex movements. This can be useful to clinicians, as well as to researchers studying multimodal communication or human–robot interactions. By making this protocol available, we hope to provide a tool that can be applied to understanding meaningful movement characteristics in human communication.

Abstract

Background Rapid automatised naming (RAN) and rapid alternating stimulus (RAS) are reliable predictors of reading disability. The underlying biology of reading disability is poorly understood. However, the high correlation among RAN, RAS and reading could be attributable to shared genetic factors that contribute to common biological mechanisms.

Objective To identify shared genetic factors that contribute to RAN and RAS performance using a multivariate approach.

Methods We conducted a multivariate genome-wide association analysis of RAN Objects, RAN Letters and RAS Letters/Numbers in a sample of 1331 Hispanic American and African–American youth. Follow-up neuroimaging genetic analysis of cortical regions associated with reading ability in an independent sample and epigenetic examination of extant data predicting tissue-specific functionality in the brain were also conducted.

Results Genome-wide significant effects were observed at rs1555839 (p=4.03×10−8) and replicated in an independent sample of 318 children of European ancestry. Epigenetic analysis and chromatin state models of the implicated 70 kb region of 10q23.31 support active transcription of the gene RNLS in the brain, which encodes a catecholamine metabolising protein. Chromatin contact maps of adult hippocampal tissue indicate a potential enhancer–promoter interaction regulating RNLS expression. Neuroimaging genetic analysis in an independent, multiethnic sample (n=690) showed that rs1555839 is associated with structural variation in the right inferior parietal lobule.

Conclusion This study provides support for a novel trait locus at chromosome 10q23.31 and proposes a potential gene–brain–behaviour relationship for targeted future functional analysis to understand underlying biological mechanisms for reading disability.

Abstract

The current study investigated the comprehension of subject and object relative clauses (RCs) in bilingual Mandarin-English children (N = 55, Mage = 7;5, SD = 1;8) and language-matched monolingual Mandarin-speaking children (N = 59, Mage = 5;4, SD = 0;7). The children completed a referent selection task that tested their comprehension of subject and object RCs, and standardised assessments of vocabulary knowledge. Results showed a very similar pattern of responding in both groups. In comparison to past studies of Cantonese, the bilingual and monolingual children both showed a significant subject-over-object RC advantage. An error analysis suggested that the children’s difficulty with object RCs reflected the tendency to interpret the sentential subject as the head noun. A subsequent corpus analysis suggested that children’s difficulty with object RCs may be in part due to distributional information favouring subject RC analyses. Individual differences analyses suggested cross-linguistic transfer from English to Mandarin in the bilingual children at the individual but not the group level, with the results indicating that comparative English-dominance makes children vulnerable to error

Abstract

Numerous studies have revealed an asymmetry tied to the perception of coronal place of articulation: participants accept a labial mispronunciation of a coronal target, but not vice versa. Whether or not this asymmetry is based on language-general properties or arises from language-specific experience has been a matter of debate. The current study suggests a bias of the first type by documenting an early, cross-linguistic asymmetry related to coronal place of articulation. Japanese and Dutch 4- and 6-month-old infants showed evidence of discrimination if they were habituated to a labial and then tested on a coronal sequence, but not vice versa. This finding has important implications for both phonological theories and infant speech perception research

Abstract

Two artificial-language learning experiments directly compared English, French, and Dutch listeners’ use of suprasegmental cues for continuous-speech segmentation. In both experiments, listeners heard unbroken sequences of consonant-vowel syllables, composed of recurring three- and four-syllable “words.” These words were demarcated by(a) no cue other than transitional probabilities induced by their recurrence, (b) a consistent left-edge cue, or (c) a consistent right-edge cue. Experiment 1 examined a vowel lengthening cue. All three listener groups benefited from this cue in right-edge position; none benefited from it in left-edge position. Experiment 2 examined a pitch-movement cue. English listeners used this cue in left-edge position, French listeners used it in right-edge position, and Dutch listeners used it in both positions. These findings are interpreted as evidence of both language-universal and language-specific effects. Final lengthening is a language-universal effect expressing a more general (non-linguistic) mechanism. Pitch movement expresses prominence which has characteristically different placements across languages: typically at right edges in French, but at left edges in English and Dutch. Finally, stress realization in English versus Dutch encourages greater attention to suprasegmental variation by Dutch than by English listeners, allowing Dutch listeners to benefit from an informative pitch-movement cue even in an uncharacteristic position.

Abstract

A dominant hypothesis in empirical research on the evolution of language is the following: the fundamental difference between animal and human communication systems is captured by the distinction between regular and more complex non-regular grammars. Studies reporting successful artificial grammar learning of nested recursive structures and imaging studies of the same have methodological shortcomings since they typically allow explicit problem solving strategies and this has been shown to account for the learning effect in subsequent behavioral studies. The present study overcomes these shortcomings by using subtle violations of agreement structure in a preference classification task. In contrast to the studies conducted so far, we use an implicit learning paradigm, allowing the time needed for both abstraction processes and consolidation to take place. Our results demonstrate robust implicit learning of recursively embedded structures (context-free grammar) and recursive structures with cross-dependencies (context-sensitive grammar) in an artificial grammar learning task spanning 9 days. Keywords: Implicit artificial grammar learning; centre embedded; cross-dependency; implicit learning; context-sensitive grammar; context-free grammar; regular grammar; non-regular grammar

Abstract

A commonly held assumption in cognitive neuroscience is that, because measures of human brain function are closer to underlying biology than distal indices of behavior/cognition, they hold more promise for uncovering genetic pathways. Supporting this view is an influential fMRI-based study of sentence reading/listening by Pinel et al. (2012), who reported that common DNA variants in specific candidate genes were associated with altered neural activation in language-related regions of healthy individuals that carried them. In particular, different single-nucleotide polymorphisms (SNPs) of FOXP2 correlated with variation in task-based activation in left inferior frontal and precentral gyri, whereas a SNP at the KIAA0319/TTRAP/THEM2 locus was associated with variable functional asymmetry of the superior temporal sulcus. Here, we directly test each claim using a closely matched neuroimaging genetics approach in independent cohorts comprising 427 participants, four times larger than the original study of 94 participants. Despite demonstrating power to detect associations with substantially smaller effect sizes than those of the original report, we do not replicate any of the reported associations. Moreover, formal Bayesian analyses reveal substantial to strong evidence in support of the null hypothesis (no effect). We highlight key aspects of the original investigation, common to functional neuroimaging genetics studies, which could have yielded elevated false-positive rates. Genetic accounts of individual differences in cognitive functional neuroimaging are likely to be as complex as behavioral/cognitive tests, involving many common genetic variants, each of tiny effect. Reliable identification of true biological signals requires large sample sizes, power calculations, and validation in independent cohorts with equivalent paradigms.

SIGNIFICANCE STATEMENT A pervasive idea in neuroscience is that neuroimaging-based measures of brain function, being closer to underlying neurobiology, are more amenable for uncovering links to genetics. This is a core assumption of prominent studies that associate common DNA variants with altered activations in task-based fMRI, despite using samples (10–100 people) that lack power for detecting the tiny effect sizes typical of genetically complex traits. Here, we test central findings from one of the most influential prior studies. Using matching paradigms and substantially larger samples, coupled to power calculations and formal Bayesian statistics, our data strongly refute the original findings. We demonstrate that neuroimaging genetics with task-based fMRI should be subject to the same rigorous standards as studies of other complex traits.

Abstract

The contribution of rare and low-frequency variants to human traits is largely unexplored. Here we describe insights from sequencing whole genomes (low read depth, 7×) or exomes (high read depth, 80×) of nearly 10,000 individuals from population-based and disease collections. In extensively phenotyped cohorts we characterize over 24 million novel sequence variants, generate a highly accurate imputation reference panel and identify novel alleles associated with levels of triglycerides (APOB), adiponectin (ADIPOQ) and low-density lipoprotein cholesterol (LDLR and RGAG1) from single-marker and rare variant aggregation tests. We describe population structure and functional annotation of rare and low-frequency variants, use the data to estimate the benefits of sequencing for association studies, and summarize lessons from disease-specific collections. Finally, we make available an extensive resource, including individual-level genetic and phenotypic data and web-based tools to facilitate the exploration of association results.

Abstract

This paper describes a new flexible delexicalization method based on glottal excited parametric speech synthesis scheme. The system utilizes inverse filtered glottal flow and all-pole modelling of the vocal tract. The method provides a possibility to retain and manipulate all relevant prosodic features of any kind of speech. Most importantly, the features include voice quality, which has not been properly modeled in earlier delexicalization methods. The functionality of the new method was tested in a prosodic tagging experiment aimed at providing word prominence data for a text-to-speech synthesis system. The experiment confirmed the usefulness of the method and further corroborated earlier evidence that linguistic factors influence the perception of prosodic prominence.

Abstract

How does the brain respond to statements that clash with a person's value system? We recorded event-related brain potentials while respondents from contrasting political-ethical backgrounds completed an attitude survey on drugs, medical ethics, social conduct, and other issues. Our results show that value-based disagreement is unlocked by language extremely rapidly, within 200 to 250 ms after the first word that indicates a clash with the reader's value system (e.g., "I think euthanasia is an acceptable/unacceptable…"). Furthermore, strong disagreement rapidly influences the ongoing analysis of meaning, which indicates that even very early processes in language comprehension are sensitive to a person's value system. Our results testify to rapid reciprocal links between neural systems for language and for valuation.

Abstract

In this chapter, I review my EEG research on comprehending sentences in context from a pragmatics-oriented perspective. The review is organized around four questions: (1) When and how do extra-sentential factors such as the prior text, identity of the speaker, or value system of the comprehender affect the incremental sentence interpretation processes indexed by the so-called N400 component of the ERP? (2) When and how do people identify the referents for expressions such as “he” or “the review”, and how do referential processes interact with sense and syntax? (3) How directly pragmatic are the interpretation-relevant ERP effects reported here? (4) Do readers and listeners anticipate upcoming information? One important claim developed in the chapter is that the well-known N400 component, although often associated with ‘semantic integration’, only indirectly reflects the sense-making involved in structure-sensitive dynamic composition of the type studied in semantics and pragmatics. According to the multiple-cause intensified retrieval (MIR) account -- essentially an extension of the memory retrieval account proposed by Kutas and colleagues -- the amplitude of the word-elicited N400 reflects the computational resources used in retrieving the relatively invariant coded meaning stored in semantic long-term memory for, and made available by, the word at hand. Such retrieval becomes more resource-intensive when the coded meanings cued by this word do not match with expectations raised by the relevant interpretive context, but also when certain other relevance signals, such as strong affective connotation or a marked delivery, indicate the need for deeper processing. The most important consequence of this account is that pragmatic modulations of the N400 come about not because the N400 at hand directly reflects a rich compositional-semantic and/or Gricean analysis to make sense of the word’s coded meaning in this particular context, but simply because the semantic and pragmatic implications of the preceding words have already been computed, and now define a less or more helpful interpretive background within which to retrieve coded meaning for the critical word.

Abstract

Speech prosody, the variation in sentence melody and rhythm, plays a crucial role in sentence comprehension. Specifically, changes in intonational pitch along a sentence can affect our understanding of who did what to whom. To date, it remains unclear how the brain processes this particular use of intonation and which brain regions are involved. In particular, one central matter of debate concerns the lateralisation of intonation processing. To study the role of intonation in sentence comprehension, we designed a functional MRI experiment in which participants listened to spoken sentences. Critically, the interpretation of these sentences depended on either intonational or grammatical cues. Our results
showed stronger functional activity in the left inferior frontal gyrus (IFG) when the intonational cue was crucial for sentence comprehension compared to when it was not. When instead a grammatical cue was crucial for sentence comprehension, we found involvement of an overlapping region in the left IFG, as well as in a posterior temporal
region. A further analysis revealed that the lateralisation of intonation processing depends on its role in syntactic processing: activity in the IFG was lateralised to the left hemisphere when intonation was the only source of information to comprehend the sentence. In contrast, activity in the IFG was right-lateralised when intonation did not contribute to sentence comprehension. Together, these results emphasise the key role of the left IFG in sentence comprehension, showing the importance of this region when intonation
establishes sentence structure. Furthermore, our results provide evidence for the theory
that the lateralisation of prosodic processing is modulated by its linguistic role.

Abstract

In synaesthetes specific sensory stimuli (e.g., black letters) elicit additional experiences (e.g. colour). Synaesthesia is highly prevalent among individuals with autism spectrum disorder but the mechanisms of this co-occurrence are not clear. We hypothesized autism and synaesthesia share atypical sensory sensitivity and perception. We assessed autistic traits, sensory sensitivity, and visual perception in two synaesthete populations. In Study 1, synaesthetes (N=79, of different types) scored higher than non-synaesthetes (N=76) on the Attention-to-detail and Social skills subscales of the Autism Spectrum Quotient indexing autistic traits, and on the Glasgow Sensory Questionnaire indexing sensory hypersensitivity and hyposensitivity which frequently occur in autism. Synaesthetes performed two local/global visual tasks because individuals with autism typically show a bias toward detail processing. In synaesthetes, elevated motion coherence thresholds suggested reduced global motion perception and higher accuracy on an embedded figures task suggested enhanced local perception. In Study 2 sequence-space synaesthetes (N=18) completed the same tasks. Questionnaire and embedded figures results qualitatively resembled Study 1 results but no significant group differences with non-synaesthetes (N=20) were obtained. Unexpectedly, sequence-space synaesthetes had reduced motion coherence thresholds. Altogether, our studies suggest atypical sensory sensitivity and a bias towards detail processing are shared features of synaesthesia and autism spectrum disorder.

Abstract

In simultaneous interpreting, speech comprehension and production processes have to be coordinated in close temporal proximity. To examine the coordination, Dutch-English bilingual participants were presented with narrative fragments recorded in English at speech rates varying from 100 to 200 words per minute and they were asked to translate the fragments into Dutch (interpreting) or repeat them in English (shadowing). Interpreting yielded more errors than shadowing at every speech rate, and increasing speech rate had a stronger negative effect on interpreting than on shadowing. To understand the differential effect of speech rate, a computational model was created of sub-lexical and lexical processes in comprehension and production. Computer simulations revealed that the empirical findings could be captured by assuming a bottleneck preventing simultaneous lexical selection in production and comprehension. To conclude, our empirical and modelling results suggest the existence of a lexical bottleneck that limits the translation of narratives at high speed.

Abstract

Common genetic variants have been identified for adult height, but not much is known about the genetics of skeletal growth in early life. To identify common genetic variants that influence fetal skeletal growth, we meta-analyzed 22 genome-wide association studies (Stage 1; N = 28 459). We identified seven independent top single nucleotide polymorphisms (SNPs) (P < 1 × 10(-6)) for birth length, of which three were novel and four were in or near loci known to be associated with adult height (LCORL, PTCH1, GPR126 and HMGA2). The three novel SNPs were followed-up in nine replication studies (Stage 2; N = 11 995), with rs905938 in DC-STAMP domain containing 2 (DCST2) genome-wide significantly associated with birth length in a joint analysis (Stages 1 + 2; β = 0.046, SE = 0.008, P = 2.46 × 10(-8), explained variance = 0.05%). Rs905938 was also associated with infant length (N = 28 228; P = 5.54 × 10(-4)) and adult height (N = 127 513; P = 1.45 × 10(-5)). DCST2 is a DC-STAMP-like protein family member and DC-STAMP is an osteoclast cell-fusion regulator. Polygenic scores based on 180 SNPs previously associated with human adult stature explained 0.13% of variance in birth length. The same SNPs explained 2.95% of the variance of infant length. Of the 180 known adult height loci, 11 were genome-wide significantly associated with infant length (SF3B4, LCORL, SPAG17, C6orf173, PTCH1, GDF5, ZNFX1, HHIP, ACAN, HLA locus and HMGA2). This study highlights that common variation in DCST2 influences variation in early growth and adult height.

Abstract

The syntactic structure of main and subordinate clauses is determined to a considerable extent by verb biases. For example, some English and Dutch ditransitive verbs have a preference for the prepositional object dative, whereas others are typically used with the double object dative. In this study, we compare the effect of these biases on structure selection in (S)VO and (S)OV dative clauses in the Corpus of Spoken Dutch (CGN). This comparison allowed us to make inferences about the size of the advance planning scope during spontaneous speaking: If the verb is an obligatory component of clause-level advance planning scope, as is claimed by the hypothesis of hierarchical incrementality, then biases should exert their influence on structure choices, regardless of early (VO) or late (OV) position of the verb in the clause. Conversely, if planning proceeds in a piecemeal fashion, strictly guided by lexical availability, as claimed by linear incrementality, then the verb and its associated biases can only influence structure choices in VO sentences. We tested these predictions by analyzing structure choices in the CGN, using mixed logit models. Our results support a combination of linear and hierarchical incrementality, showing a significant influence of verb bias on structure choices in VO, and a weaker (but still significant) effect in OV clauses

Abstract

A not uncommon assumption in psycholinguistics is that the N400 directly indexes high-level semantic integration, the compositional, word-driven construction of sentence- and discourse-level meaning in some language-relevant unification space. The various discourse- and speaker-dependent modulations of the N400 uncovered by us and others are often taken to support this 'compositional integration' position. In my talk, I will argue that these N400 modulations are probably better interpreted as only indirectly reflecting compositional sense-making. The account that I will advance for these N400 effects is a variant of the classic Kutas and Federmeier (2002, TICS) memory retrieval account in which context effects on the word-elicited N400 are taken to reflect contextual priming of LTM access. It differs from the latter in making more explicit that the contextual cues that prime access to a word's meaning in LTM can range from very simple (e.g., a single concept) to very complex ones (e.g., a structured representation of the current discourse). Furthermore, it incorporates the possibility, suggested by recent N400 findings, that semantic retrieval can also be intensified in response to certain ‘relevance signals’, such as strong value-relevance, or a marked delivery (linguistic focus, uncommon choice of words, etc). In all, the perspective I'll draw is that in the context of discourse-level language processing, N400 effects reflect an 'overlay of technologies', with the construction of discourse-level representations riding on top of more ancient sense-making technology.

Abstract

This study explores the effect of feedback with hints on students’ recall of words. In three classroom experiments, high school students individually practiced vocabulary words through computerized retrieval practice with either standard show-answer feedback (display of answer) or hints feedback after incorrect responses. Hints feedback gave students a second chance to find the correct response using orthographic (Experiment 1), mnemonic (Experiment 2), or cross-language hints (Experiment 3). During practice, hints led to a shift of practice time from further repetitions to longer feedback processing but did not reduce (repeated) errors. There was no effect of feedback on later recall except when the hints from practice were also available on the test, indicating limited transfer of practice with hints to later recall without hints (in Experiments 1 and 2). Overall, hints feedback was not preferable over show-answer feedback. The common notion that hints are beneficial may not hold when the total practice time is limited.

Abstract

The temporal properties of semantic and phonological processes in speech production were investigated in a new experimental paradigm using movement-related brain potentials. The main experimental task was picture naming. In addition, a 2-choice reaction go/no-go procedure was included, involving a semantic and a phonological categorization of the picture name. Lateralized readiness potentials (LRPs) were derived to test whether semantic and phonological information activated motor processes at separate moments in time. An LRP was only observed on no-go trials when the semantic (not the phonological) decision determined the response hand. Varying the position of the critical phoneme in the picture name did not affect the onset of the LRP but rather influenced when the LRP began to differ on go and no-go trials and allowed the duration of phonological encoding of a word to be estimated. These results provide electrophysiological evidence for early semantic activation and later phonological encoding.