The language production system of literate adults comprises an orthographic system (used during written language production) and a phonological system (used during spoken language production). Recent psycholinguistic research has investigated possible influences of the orthographic system on the phonological system. This research has produced contrastive results, with some studies showing effects of orthography in the course of normal speech production while others failing to show such effects. In this article, we review the available evidence and consider possible explanations for the discrepancy. We then report two form-preparation experiments which aimed at testing for the effects of orthography in spoken word-production. Our results provide clear evidence that the orthographic properties of the words do not influence their spoken production in picture naming. We discuss this finding in relation to psycholinguistic and neuropsychological investigations of the relationship between written and spoken word-production.

There is growing evidence suggesting that synchronization changes in the oscillatory neuronal dynamics in the EEG or MEG reflect the transient coupling and uncoupling of functional networks related to different aspects of language comprehension. In this work, we examine how sentence-level syntactic unification operations are reflected in the oscillatory dynamics of the MEG. Participants read sentences that were either correct, contained a word category violation, or were constituted of random word sequences devoid of syntactic structure. A time-frequency analysis of MEG power changes revealed three types of effects. The first type of effect was related to the detection of a (word category) violation in a syntactically structured sentence, and was found in the alpha and gamma frequency bands. A second type of effect was maximally sensitive to the syntactic manipulations: A linear increase in beta power across the sentence was present for correct sentences, was disrupted upon the occurrence of a word category violation, and was absent in syntactically unstructured random word sequences. We therefore relate this effect to syntactic unification operations. Thirdly, we observed a linear increase in theta power across the sentence for all syntactically structured sentences. The effects are tentatively related to the building of a working memory trace of the linguistic input. In conclusion, the data seem to suggest that syntactic unification is reflected by neuronal synchronization in the lower-beta frequency band.

Functional organization of the lateral temporal cortex in humans is not well understood. We recorded blood oxygenation signals from the temporal lobes of normal volunteers using functional magnetic resonance imaging during stimulation with unstructured noise, frequency-modulated (FM) tones, reversed speech, pseudowords and words. For all conditions, subjects performed a material-nonspecific detection response when a train of stimuli began or ceased. Dorsal areas surrounding Heschl's gyrus bilaterally, particularly the planum temporale and dorsolateral superior temporal gyrus, were more strongly activated by FM tones than by noise, suggesting a role in processing simple temporally encoded auditory information. Distinct from these dorsolateral areas, regions centered in the superior temporal sulcus bilaterally were more activated by speech stimuli than by FM tones. Identical results were obtained in this region using words, pseudowords and reversed speech, suggesting that the speech-tones activation difference is due to acoustic rather than linguistic factors. In contrast, previous comparisons between word and nonword speech sounds showed left-lateralized activation differences in more ventral temporal and temporoparietal regions that are likely involved in processing lexical-semantic or syntactic information associated with words. The results indicate functional subdivision of the human lateral temporal cortex and provide a preliminary framework for understanding the cortical processing of speech sounds.

Recently, we introduced a new 'GLM-beamformer' technique for MEG analysis that enables accurate localisation of both phase-locked and non-phase-locked neuromagnetic effects, and their representation as statistical parametric maps (SPMs). This provides a useful framework for comparison of the full range of MEG responses with fMRI BOLD results. This paper reports a 'proof of principle' study using a simple visual paradigm (static checkerboard). The five subjects each underwent both MEG and fMRI paradigms. We demonstrate, for the first time, the presence of a sustained (DC) field in the visual cortex, and its co-localisation with the visual BOLD response. The GLM-beamformer analysis method is also used to investigate the main non-phase-locked oscillatory effects: an event-related desynchronisation (ERD) in the alpha band (8-13 Hz) and an event-related synchronisation (ERS) in the gamma band (55-70 Hz). We show, using SPMs and virtual electrode traces, the spatio-temporal covariance of these effects with the visual BOLD response. Comparisons between MEG and fMRI data sets generally focus on the relationship between the BOLD response and the transient evoked response. Here, we show that the stationary field and changes in oscillatory power are also important contributors to the BOLD response, and should be included in future studies on the relationship between neuronal activation and the haemodynamic response.

The transformation of an abstract phonological code into articulation has been hypothesized to involve the retrieval of stored syllable-sized motor plans. Accordingly, gestural scores for frequently used syllables are retrieved from memory whereas gestural scores for novel and possibly low frequency syllables are assembled on-line. The present study was designed to test this hypothesis. Participants produced disyllabic pseudowords with high frequency, low frequency and non-existent (novel) initial syllables. Behavioral results revealed slower production latencies for novel than for high frequency syllables. Event-related potentials diverged in waveform amplitudes and global topographic patterns between high frequency and low frequency/novel syllables around 170 ms before the onset of articulation. These differences indicate the recruitment of different brain networks during the production of frequent and infrequent/novel syllables, in line with the hypothesis that speakers store syllabic-sized motor programs for frequent syllables and assemble these motor plans on-line for low frequency and novel syllables.

Electroencephalography (EEG) has been instrumental in making discoveries about cognition, brain function, and dysfunction. However, where do EEG signals come from and what do they mean? The purpose of this paper is to argue that we know shockingly little about the answer to this question, to highlight what we do know, how important the answers are, and how modern neuroscience technologies that allow us to measure and manipulate neural circuits with high spatiotemporal accuracy might finally bring us some answers. Neural oscillations are perhaps the best feature of EEG to use as anchors because oscillations are observed and are studied at multiple spatiotemporal scales of the brain, in multiple species, and are widely implicated in cognition and in neural computations.

In recent years, studies ranging from single-unit recordings in animals to electroencephalography and magnetoencephalography studies in humans have demonstrated the pivotal role of phase synchronization in memory processes. Phase synchronization - here referring to the synchronization of oscillatory phases between different brain regions - supports both working memory and long-term memory and acts by facilitating neural communication and by promoting neural plasticity. There is evidence that processes underlying working and long-term memory might interact in the medial temporal lobe. We propose that this is accomplished by neural operations involving phase-phase and phase-amplitude synchronization. A deeper understanding of how phase synchronization supports the flexibility of and interaction between memory systems may yield new insights into the functions of phase synchronization in general.

UNLABELLED: Using a continuous listening task, we evaluated the coupling between the listener's cortical activity and the temporal envelopes of different sounds in a multitalker auditory scene using magnetoencephalography and corticovocal coherence analysis. Neuromagnetic signals were recorded from 20 right-handed healthy adult humans who listened to five different recorded stories (attended speech streams), one without any multitalker background (No noise) and four mixed with a

Neuronal oscillations are ubiquitous in the brain and may contribute to cognition in several ways: for example, by segregating information and organizing spike timing. Recent data show that delta, theta and gamma oscillations are specifically engaged by the multi-timescale, quasi-rhythmic properties of speech and can track its dynamics. We argue that they are foundational in speech and language processing, 'packaging' incoming information into units of the appropriate temporal granularity. Such stimulus-brain alignment arguably results from auditory and motor tuning throughout the evolution of speech and language and constitutes a natural model system allowing auditory research to make a unique contribution to the issue of how neural oscillatory activity affects human cognition.

Plants are continually exposed to pathogen attack but usually remain healthy because they can activate defences upon perception of microbes. However, pathogens have evolved to overcome plant immunity by delivering effectors into the plant cell to attenuate defence, resulting in disease. Recent studies suggest that some effectors may manipulate host transcription, but the specific mechanisms by which such effectors promote susceptibility remain unclear. We study the oomycete downy mildew pathogen of Arabidopsis, Hyaloperonospora arabidopsidis (Hpa), and show here that the nuclear-localized effector HaRxL44 interacts with Mediator subunit 19a (MED19a), resulting in the degradation of MED19a in a proteasome-dependent manner. The Mediator complex of approximately 25 proteins is broadly conserved in eukaryotes and mediates the interaction between transcriptional regulators and RNA polymerase II. We found MED19a to be a positive regulator of immunity against Hpa. Expression profiling experiments reveal transcriptional changes resembling jasmonic acid/ethylene (JA/ET) signalling in the presence of HaRxL44, and also 3 d after infection with Hpa. Elevated JA/ET signalling is associated with a decrease in salicylic acid (SA)-triggered immunity (SATI) in Arabidopsis plants expressing HaRxL44 and in med19a loss-of-function mutants, whereas SATI is elevated in plants overexpressing MED19a. Using a PR1::GUS reporter, we discovered that Hpa suppresses PR1 expression specifically in cells containing haustoria, into which RxLR effectors are delivered, but not in nonhaustoriated adjacent cells, which show high PR1::GUS expression levels. Thus, HaRxL44 interferes with Mediator function by degrading MED19, shifting the balance of defence transcription from SA-responsive defence to JA/ET-signalling, and enhancing susceptibility to biotrophs by attenuating SA-dependent gene expression.

The formation of cortical object representations requires the activation of cell assemblies, correlated by induced oscillatory bursts of activity > 20 Hz (induced gamma band responses; iGBRs). One marker of the functional dynamics within such cell assemblies is the suppression of iGBRs elicited by repeated stimuli. This effect is commonly interpreted as a signature of 'sharpening' processes within cell-assemblies, which are behaviourally mirrored in repetition priming effects. The present study investigates whether the sharpening of primed objects is an automatic consequence of repeated stimulus processing, or whether it depends on task demands. Participants performed either a 'living/non-living' or a 'bigger/smaller than a shoebox' classification on repeated pictures of everyday objects. We contrasted repetition-related iGBR effects after the same task was used for initial and repeated presentations (no-switch condition) with repetitions after a task-switch occurred (switch condition). Furthermore, we complemented iGBR analysis by examining other brain responses known to be modulated by repetition-related memory processes (evoked gamma oscillations and event-related potentials; ERPs). The results obtained for the 'no-switch' condition replicated previous findings of repetition suppression of iGBRs at 200-300 ms after stimulus onset. Source modelling showed that this effect was distributed over widespread cortical areas. By contrast, after a task-switch no iGBR suppression was found. We concluded that iGBRs reflect the sharpening of a cell assembly only within the same task. After a task switch the complete object representation is reactivated. The ERP (220-380 ms) revealed suppression effects independent of task demands in bilateral posterior areas and might indicate correlates of repetition priming in perceptual structures.

Do we say dog when we say hotdog? In five experiments using the implicit priming paradigm, we assessed whether nominal compounds composed of two free morphemes like sawdust or fishbowl are prepared for production at the segmental level in the same way that two-syllable monomorphemic words (e.g. bandit) are, or instead as sequences of separable words (e.g. full bowl or grey dust). The experiments demonstrated that nominal compounds are planned as a single sequence, not as two sequences. Specifically, the onset of the second component of the compound (e.g. /d/ in sawdust) did not act as a primeable starting point, although comparable onsets did when that component was an independent word (grey dust). We conclude that there may be a dog in hotdog at the morpheme level, but not when phonological segments are prepared for production.

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

The role of neuronal oscillations during language comprehension is not yet well understood. In this paper we review and reinterpret the functional roles of beta- and gamma-band oscillatory activity during language comprehension at the sentence and discourse level. We discuss the evidence in favor of a role for beta and gamma in unification (the unification hypothesis), and in light of mounting evidence that cannot be accounted for under this hypothesis, we explore an alternative proposal linking beta and gamma oscillations to maintenance and prediction (respectively) during language comprehension. Our maintenance/prediction hypothesis is able to account for most of the findings that are currently available relating beta and gamma oscillations to language comprehension, and is in good agreement with other proposals about the roles of beta and gamma in domain-general cognitive processing. In conclusion we discuss proposals for further testing and comparing the prediction and unification hypotheses.

SummaryHow natural speech is represented in the auditory cortex constitutes a major challenge for cognitive neuroscience. Although many single-unit and neuroimaging studies have yielded valuable insights about the processing of speech and matched complex sounds, the mechanisms underlying the analysis of speech dynamics in human auditory cortex remain largely unknown. Here, we show that the phase pattern of theta band (4–8 Hz) responses recorded from human auditory cortex with magnetoencephalography (MEG) reliably tracks and discriminates spoken sentences and that this discrimination ability is correlated with speech intelligibility. The findings suggest that an ∼200 ms temporal window (period of theta oscillation) segments the incoming speech signal, resetting and sliding to track speech dynamics. This hypothesized mechanism for cortical speech analysis is based on the stimulus-induced modulation of inherent cortical rhythms and provides further evidence implicating the syllable as a computational primitive for the representation of spoken language.]]>

In this paper, we show how ElectroEncephaloGraphic (EEG) and MagnetoEncephaloGraphic (MEG) data can be analyzed statistically using nonparametric techniques. Nonparametric statistical tests offer complete freedom to the user with respect to the test statistic by means of which the experimental conditions are compared. This freedom provides a straightforward way to solve the multiple comparisons problem (MCP) and it allows to incorporate biophysically motivated constraints in the test statistic, which may drastically increase the sensitivity of the statistical test. The paper is written for two audiences: (1) empirical neuroscientists looking for the most appropriate data analysis method, and (2) methodologists interested in the theoretical concepts behind nonparametric statistical tests. For the empirical neuroscientist, a large part of the paper is written in a tutorial-like fashion, enabling neuroscientists to construct their own statistical test, maximizing the sensitivity to the expected effect. And for the methodologist, it is explained why the nonparametric test is formally correct. This means that we formulate a null hypothesis (identical probability distribution in the different experimental conditions) and show that the nonparametric test controls the false alarm rate under this null hypothesis.

Developmental dyslexia is a reading disorder often characterized by reduced awareness of speech units. Whether the neural source of this phonological disorder in dyslexic readers results from the malfunctioning of the primary auditory system or damaged feedback communication between higher-order phonological regions (i.e., left inferior frontal regions) and the auditory cortex is still under dispute. Here we recorded magnetoencephalographic (MEG) signals from 20 dyslexic readers and 20 age-matched controls while they were listening to approximately 10-s-long spoken sentences. Compared to controls, dyslexic readers had (1) an impaired neural entrainment to speech in the delta band (0.5-1 Hz); (2) a reduced delta synchronization in both the right auditory cortex and the left inferior frontal gyrus; and (3) an impaired feedforward functional coupling between neural oscillations in the right auditory cortex and the left inferior frontal regions. This shows that during speech listening, individuals with developmental dyslexia present reduced neural synchrony to low-frequency speech oscillations in primary auditory regions that hinders higher-order speech processing steps. The present findings, thus, strengthen proposals assuming that improper low-frequency acoustic entrainment affects speech sampling. This low speech-brain synchronization has the strong potential to cause severe consequences for both phonological and reading skills. Interestingly, the reduced speech-brain synchronization in dyslexic readers compared to normal readers (and its higher-order consequences across the speech processing network) appears preserved through the development from childhood to adulthood. Thus, the evaluation of speech-brain synchronization could possibly serve as a diagnostic tool for early detection of children at risk of dyslexia. Hum Brain Mapp 37:2767-2783, 2016. (c) 2016 Wiley Periodicals, Inc.

This paper presents

In Mandarin Chinese, speakers benefit from fore-knowledge of what the first syllable but not of what the first phonemic segment of a disyllabic word will be (Chen, Chen, & Dell, 2002), contrasting with findings in English, Dutch, and other Indo-European languages, and challenging the generality of current theories of word production. In this article, we extend the evidence for the language difference by showing that failure to prepare onsets in Mandarin (Experiment 1) applies even to simple monosyllables (Experiments 2-4), and confirm the contrast with English for comparable materials (Experiments 5 and 6). We also provide new evidence that Mandarin speakers do reliably prepare tonally unspecified phonological syllables (Experiment 7). To account for these patterns, we propose a language general proximate units principle whereby intentional preparation for speech as well as phonological-lexical coordination are grounded at the first phonological level below the word at which explicit unit selection occurs. The language difference arises because syllables are proximate units in Mandarin Chinese, whereas segments are proximate in English and other Indo-European languages. The proximate units perspective reconciles the aspiration toward a language general account of word production with the reality of substantial cross-linguistic differences.

A growing body of evidence shows that ongoing oscillations in auditory cortex modulate their phase to match the rhythm of temporally regular acoustic stimuli, increasing sensitivity to relevant environmental cues and improving detection accuracy. In the current study, we test the hypothesis that nonsensory information provided by linguistic content enhances phase-locked responses to intelligible speech in the human brain. Sixteen adults listened to meaningful sentences while we recorded neural activity using magnetoencephalography. Stimuli were processed using a noise-vocoding technique to vary intelligibility while keeping the temporal acoustic envelope consistent. We show that the acoustic envelopes of sentences contain most power between 4 and 7 Hz and that it is in this frequency band that phase locking between neural activity and envelopes is strongest. Bilateral oscillatory neural activity phase-locked to unintelligible speech, but this cerebro-acoustic phase locking was enhanced when speech was intelligible. This enhanced phase locking was left lateralized and localized to left temporal cortex. Together, our results demonstrate that entrainment to connected speech does not only depend on acoustic characteristics, but is also affected by listeners ability to extract linguistic information. This suggests a biological framework for speech comprehension in which acoustic and linguistic cues reciprocally aid in stimulus prediction.

Recent psychophysics data suggest that speech perception is not limited by the capacity of the auditory system to encode fast acoustic variations through neural gamma activity, but rather by the time given to the brain to decode them. Whether the decoding process is bounded by the capacity of theta rhythm to follow syllabic rhythms in speech, or constrained by a more endogenous top-down mechanism, e.g., involving beta activity, is unknown. We addressed the dynamics of auditory decoding in speech comprehension by challenging syllable tracking and speech decoding using comprehensible and incomprehensible time-compressed auditory sentences. We recorded EEGs in human participants and found that neural activity in both theta and gamma ranges was sensitive to syllabic rate. Phase patterns of slow neural activity consistently followed the syllabic rate (4-14 Hz), even when this rate went beyond the classical theta range (4-8 Hz). The power of theta activity increased linearly with syllabic rate but showed no sensitivity to comprehension. Conversely, the power of beta (14-21 Hz) activity was insensitive to the syllabic rate, yet reflected comprehension on a single-trial basis. We found different long-range dynamics for theta and beta activity, with beta activity building up in time while more contextual information becomes available. This is consistent with the roles of theta and beta activity in stimulus-driven versus endogenous mechanisms. These data show that speech comprehension is constrained by concurrent stimulus-driven theta and low-gamma activity, and by endogenous beta activity, but not primarily by the capacity of theta activity to track the syllabic rhythm.SIGNIFICANCE STATEMENT Speech comprehension partly depends on the ability of the auditory cortex to track syllable boundaries with theta-range neural oscillations. The reason comprehension drops when speech is accelerated could hence be because theta oscillations can no longer follow the syllabic rate. Here, we presented subjects with comprehensible and incomprehensible accelerated speech, and show that neural phase patterns in the theta band consistently reflect the syllabic rate, even when speech becomes too fast to be intelligible. The drop in comprehension, however, is signaled by a significant decrease in the power of low-beta oscillations (14-21 Hz). These data suggest that speech comprehension is not limited by the capacity of theta oscillations to adapt to syllabic rate, but by an endogenous decoding process.

Spoken sentence comprehension relies on rapid and effortless temporal integration of speech units displayed at different rates. Temporal integration refers to how chunks of information perceived at different time scales are linked together by the listener in mapping speech sounds onto meaning. The neural implementation of this integration remains unclear. This study explores the role of short and long windows of integration in accessing meaning from long samples of speech. In a cross-linguistic study, we explore the time course of oscillatory brain activity between 1 and 100 Hz, recorded using EEG, during the processing of native and foreign languages. We compare oscillatory responses in a group of Italian and Spanish native speakers while they attentively listen to Italian, Japanese, and Spanish utterances, played either forward or backward. The results show that both groups of participants display a significant increase in gamma band power (55-75 Hz) only when they listen to their native language played forward. The increase in gamma power starts around 1000 msec after the onset of the utterance and decreases by its end, resembling the time course of access to meaning during speech perception. In contrast, changes in low-frequency power show similar patterns for both native and foreign languages. We propose that gamma band power reflects a temporal binding phenomenon concerning the coordination of neural assemblies involved in accessing meaning of long samples of speech.

Description of mapping methods using spherical splines, both to interpolate scalp potentials (SPs), and to approximate scalp current densities (SCDs). Compared to a previously published method using thin plate splines, the advantages are a very simple derivation of the SCD approximation, faster computing times, and greater accuracy in areas with few electrodes.

Developments in technologic and analytical procedures applied to the study of brain electrical activity have intensified interest in this modality as a means of examining brain function. The impact of these new developments on traditional methods of acquiring and analyzing electroencephalographic activity requires evaluation. Ultimately, the integration of the old with the new must result in an accepted standardized methodology to be used in these investigations. In this paper, basic procedures and recent developments involved in the recording and analysis of brain electrical activity are discussed and recommendations are made, with emphasis on psychophysiological applications of these procedures.

Auditory cortical oscillations have been proposed to play an important role in speech perception. It is suggested that the brain may take temporal

Do speakers of all languages use segmental speech sounds when they produce words? Existing models of language production generally assume a mental representation of individual segmental units, or phonemes, but the bulk of evidence comes from speakers of European languages in which the orthographic system codes explicitly for speech sounds. By contrast, in languages with nonalphabetical scripts, such as Mandarin Chinese, individual speech sounds are not orthographically represented, raising the possibility that speakers of these languages do not use phonemes as fundamental processing units. We used event-related potentials (ERPs) combined with behavioral measurement to investigate the role of phonemes in Mandarin production. Mandarin native speakers named colored line drawings of objects using color adjective-noun phrases; color and object name either shared the initial phoneme or were phonologically unrelated. Whereas naming latencies were unaffected by phoneme repetition, ERP responses were modulated from 200 ms after picture onset. Our ERP findings thus provide strong support for the claim that phonemic segments constitute fundamental units of phonological encoding even for speakers of languages that do not encode such units orthographically.

Lexical access in speaking consists of two major steps: lemma retrieval and word-form encoding. In Roelofs (Roelofs, A. 1992a. Cognition 42. 107-142; Roelofs. A. 1993. Cognition 47, 59-87.), I described a model of lemma retrieval. The present paper extends this work by presenting a comprehensive model of the second access step, word-form encoding. The model is called WEAVER (Word-form Encoding by Activation and VERification). Unlike other models of word-form generation, WEAVER is able to provide accounts of response time data, particularly from the picture-word interference paradigm and the implicit priming paradigm. Its key features are (1) retrieval by spreading activation, (2) verification of activated information by a production rule, (3) a rightward incremental construction of phonological representations using a principle of active syllabification, syllables are constructed on the fly rather than stored with lexical items, (4) active competitive selection of syllabic motor programs using a mathematical formalism that generates response times and (5) the association of phonological speech errors with the selection of syllabic motor programs due to the failure of verification.

Neural oscillations at different frequencies have recently been related to a wide range of basic and higher cognitive processes. One possible role of oscillatory activity is to assure the maintenance of information in working memory (WM). Here we review the possibility that rhythmic activity at theta, alpha, and gamma frequencies serve distinct functional roles during WM maintenance. Specifically, we propose that gamma-band oscillations are generically involved in the maintenance of WM information. By contrast, alpha-band activity reflects the active inhibition of task-irrelevant information, whereas theta-band oscillations underlie the organization of sequentially ordered WM items. Finally, we address the role of cross-frequency coupling (CFC) in enabling alpha-gamma and theta-gamma codes for distinct WM information.

AbstractCognitive functions are organized in distributed, overlapping, and interacting brain networks. Investigation of those large-scale brain networks is a major task in neuroimaging research.Here, we introduce a novel combination of functional and anatomical connectivity to study the network topology subserving a cognitive function of interest. (i) In a given network, direct interactions between network nodes are identified by analyzing functional MRI time series with the multivariate method of directed partial correlation (dPC). This method provides important improvements over shortcomings that are typical for ordinary (partial) correlation techniques. (ii) For directly interacting pairs of nodes, a region-to-region probabilistic fiber tracking on diffusion tensor imaging data is performed to identify the most probable anatomical white matter fiber tracts mediating the functional interactions. This combined approach is applied to the language domain to investigate the network topology of two levels of auditory comprehension: lower-level speech perception (i.e., phonological processing) and higher-level speech recognition (i.e., semantic processing).For both processing levels, dPC analyses revealed the functional network topology and identified central network nodes by the number of direct interactions with other nodes. Tractography showed that these interactions are mediated by distinct ventral (via the extreme capsule) and dorsal (via the arcuate/superior longitudinal fascicle fiber system) long- and short-distance association tracts as well as commissural fibers.Our findings demonstrate how both processing routines are segregated in the brain on a large-scale network level. Combining dPC with probabilistic tractography is a promising approach to unveil how cognitive functions emerge through interaction of functionally interacting and anatomically interconnected brain regions.]]>

Reading aloud is faster when targets (e.g., PAIR) are preceded by visually masked primes sharing just the onset (e.g., pole) compared to all different primes (e.g., take). This effect is known as the masked onset priming effect (MOPE). One crucial feature of this effect is its presumed non-lexical basis. This aspect of the MOPE is tested in the current study. Dutch participants named pictures having bisyllabic names, which were preceded by visually masked primes. Picture naming was facilitated by first-segment but not last-segment primes, and by first-syllable as well as last-syllable primes. Whole-word primes with first or last segment overlap slowed down picture naming latencies significantly. The first-segment priming effect (i.e., MOPE) cannot be accounted for by non-lexical response competition since pictures cannot be named via the non-lexical route. Instead, the effects obtained in this study can be accommodated by a speech-planning account of the MOPE.

Cognition results from interactions among functionally specialized but widely distributed brain regions; however, neuroscience has so far largely focused on characterizing the function of individual brain regions and neurons therein. Here we discuss recent studies that have instead investigated the interactions between brain regions during cognitive processes by assessing correlations between neuronal oscillations in different regions of the primate cerebral cortex. These studies have opened a new window onto the large-scale circuit mechanisms underlying sensorimotor decision-making and top-down attention. We propose that frequency-specific neuronal correlations in large-scale cortical networks may be 'fingerprints' of canonical neuronal computations underlying cognitive processes.

Because context has a robust influence on the processing of subsequent words, the idea that readers and listeners predict upcoming words has attracted research attention, but prediction has fallen in and out of favor as a likely factor in normal comprehension. We note that the common sense of this word includes both benefits for confirmed predictions and costs for disconfirmed predictions. The N400 component of the event-related potential (ERP) reliably indexes the benefits of semantic context Evidence that the N400 is sensitive to the other half of prediction - a cost for failure - is largely absent from the literature. This raises the possibility that "prediction" is not a good description of what comprehenders do. However, it need not be the case that the benefits and costs of prediction are evident in a single ERP component Research outside of language processing indicates that late positive components of the ERP are very sensitive to disconfirmed predictions. We review late positive components elicited by words that are potentially more or less predictable from preceding sentence context. This survey suggests that late positive responses to unexpected words are fairly common, but that these consist of two distinct components with different scalp topographies, one associated with semantically incongruent words and one associated with congruent words. We conclude with a discussion of the possible cognitive correlates of these distinct late positivities and their relationships with more thoroughly characterized ERP components, namely the P300, P600 response to syntactic errors, and the "old/new effect" in studies of recognition memory. (C) 2011 Elsevier B.V.

It is widely acknowledged in Germanic languages that segments are the primary planning units at the phonological encoding stage of spoken word production. Mixed results, however, have been found in Chinese, and it is still unclear what roles syllables and segments play in planning Chinese spoken word production. In the current study, participants were asked to first prepare and later produce disyllabic Mandarin words upon picture prompts and a response cue while electroencephalogram (EEG) signals were recorded. Each two consecutive pictures implicitly formed a pair of prime and target, whose names shared the same word-initial atonal syllable or the same word-initial segments, or were unrelated in the control conditions. Only syllable repetition induced significant effects on event-related brain potentials (ERPs) after target onset: a widely distributed positivity in the 200- to 400-ms interval and an anterior positivity in the 400- to 600-ms interval. We interpret these to reflect syllable-size representations at the phonological encoding and phonetic encoding stages. Our results provide the first electrophysiological evidence for the distinct role of syllables in producing Mandarin spoken words, supporting a language specificity hypothesis about the primary phonological units in spoken word production.

The central problem for cognitive neuroscience is to describe how cognitive processes arise from brain processes. This review summarizes the recent evidence that synchronous neural oscillations reveal much about the origin and nature of cognitive processes such as memory, attention and consciousness. Memory processes are most closely related to theta and gamma rhythms, whereas attention seems closely associated with alpha and gamma rhythms. Conscious awareness may arise from synchronous neural oscillations occurring globally throughout the brain rather than from the locally synchronous oscillations that occur when a sensory area encodes a stimulus. These associations between the dynamics of the brain and cognitive processes indicate progress towards a unified theory of brain and cognition.

Four experiments investigated the role of the syllable in Chinese spoken word production. Chen, Chen and Ferrand (2003) reported a syllable priming effect when primes and targets shared the first syllable using a masked priming paradigm in Chinese. Our Experiment 1 was a direct replication of Chen et al.'s (2003) Experiment 3 employing CV (e.g., ,/ba2.ying2/, strike camp) and CVG (e.g., ,/bai2.shou3/, white haired) syllable types. Experiment 2 tested the syllable priming effect using different syllable types: e.g., CV (,/qi4.qiu2/, balloon) and CVN (,/qing1.ting2/, dragonfly). Experiment 3 investigated this issue further using line drawings of common objects as targets that were preceded either by a CV (e.g., ,/qi3/, attempt), or a CVN (e.g., ,/qing2/, affection) prime. Experiment 4 further examined the priming effect by a comparison between CV or CVN priming and an unrelated priming condition using CV-NX (e.g., ,/mi2.ni3/, mini) and CVN-CX (e.g., ,/min2.ju1/, dwellings) as target words. These four experiments consistently found that CV targets were named faster when preceded by CV primes than when they were preceded by CVG, CVN or unrelated primes, whereas CVG or CVN targets showed the reverse pattern. These results indicate that the priming effect critically depends on the match between the structure of the prime and that of the first syllable of the target. The effect obtained in this study was consistent across different stimuli and different tasks (word and picture naming), and provides more conclusive and consistent data regarding the role of the syllable in Chinese speech production.

Established linguistic theoretical frameworks propose that alphabetic language speakers use phonemes as phonological encoding units during speech production whereas Mandarin Chinese speakers use syllables. This framework was challenged by recent neural evidence of facilitation induced by overlapping initial phonemes, raising the possibility that phonemes also contribute to the phonological encoding process in Chinese. However, there is no evidence of non-initial phoneme involvement in Chinese phonological encoding among representative Chinese speakers, rendering the functional role of phonemes in spoken Chinese controversial. Here, we addressed this issue by systematically investigating the word-initial and non-initial phoneme repetition effect on the electrophysiological signal using a picture-naming priming task in which native Chinese speakers produced disyllabic word pairs. We found that overlapping phonemes in both the initial and non-initial position evoked more positive ERPs in the 180- to 300-ms interval, indicating position-invariant repetition facilitation effect during phonological encoding. Our findings thus revealed the fundamental role of phonemes as independent phonological encoding units in Mandarin Chinese.

Speaking involves stages of conceptual preparation, lemma selection, word-form encoding and articulation. Furthermore, process of word-form encoding can be divided into morphological encoding process, phonological encoding process and phonetic encoding. What is the function unit at the stage of word-form encoding remains a controversial issue in speech production theories. The present study investigated syllable and segments effects at the stages of phonological encoding, phonetic encoding, and articulation in Mandarin spoken word production. Using Picture-Word Interference (PWI) Paradigm, we compared the effects generated in immediately naming (experiment 1), delayed naming (experiment 2), and delayed naming combined with articulation suppression (experiment 3) tasks. Eighteen black and white line drawings were applied as stimuli, and their names were monosyllabic words. Each target picture was paired with four distractor words: A CVC-related (C: Consonant, V: Vowel) distractor word was chosen that shared a syllable which always differed in tone with the picture name (i.e.,羊 /yang2/ as target name -央/yang1/ as distractor word). A CV-related distractor word was chosen that shared the onset consonant and the core vowel with the picture name (i.e., 羊/yang2/-药/yao4/). A VC-related distractor was chosen that shared the rhymes with the picture name (i.e., 羊/yang2/-让/rang4/). An unrelated distractor was selected that stood in no obvious semantic, phonological or orthographic relation with the picture name. We found syllable and segments facilitation effects in immediate naming, whereas syllable and segments inhibition effects in a delayed naming and a combination task of delayed naming and articulation suppression. An immediate naming involves stages of phonological encoding, phonetic encoding, and articulation, a delayed naming involves articulation only, while a combination task of delayed naming and articulation suppression involves phonetic encoding and articulation processes. By comparing these effects among three tasks, we suggest that syllable and segments facilitation effects localized at the stage of phonological encoding, whereas syllable and segments inhibition effects localized at the stage of phonetic encoding and (or) articulation. These findings indicated that syllable plays a more important role in phonological encoding whereas segments play their roles in phonetic encoding and articulation for motor programming. Our findings provide support for Proximate Unit Principle and the assumption of independence of premotor- (phonological encoding) and motor stages (phonetic encoding and articulation).

The present paper addresses the syllable’s role in language production. Firstly the concept of syllable is introduced. Secondly the syllable’s position in language production theories is discussed. Then the syllable’s studies are summarized from the following two aspects: (1) the influential experimental paradigms, and (2) three important issues in this field. The main experimental paradigms include masked priming, repetition priming, implicit priming and picture-word interference paradigms. The main important issues are: (1) whether syllable is a functional unit in language production or not, (2) how syllable play important role in speech production, and (3) where is syllable priming effect. Finally, corresponding to the unique characteristics of syllable in Chinese, the syllable’s studies in Chinese word production and the perspectives of further research are discussed

Languages may differ regarding the primary mental unit of phonological encoding in spoken production, with models of speakers of Indo-European languages generally assuming a central role for phonemes, but spoken Chinese production potentially attributing a more prominent role to syllables. In the present study, native Mandarin Chinese speakers named objects that were preceded by briefly presented and masked prime words, which were form related and either matched or mismatched concerning their syllabic structure, or were unrelated. Behavioral results showed a previously reported interaction between prime and target syllable type. Concurrently recorded EEG also exhibited this interaction and further revealed that syllable overlap modulated ERPs mainly in the time window of 300-400 ms after picture onset. By contrast, phonemic overlap modulated ERPs from 500 ms to 600 ms. This pattern might suggest that speakers retrieved syllables before phonemes and strengthens the claim that for Chinese individuals syllables constitute primary functional representations (

The effects of word frequency (WF) and syllable frequency (SF) are well-established phenomena in domain such as spoken production in alphabetic languages. Chinese, as a non-alphabetic language, presents unique lexical and phonological properties in speech production. For example, the proximate unit of phonological encoding is syllable in Chinese but segments in Dutch, French or English. The present study investigated the effects of WF and SF, and their interaction in Chinese written and spoken production. Significant facilitatory WF and SF effects were observed in spoken as well as in written production. The SF effect in writing indicated that phonological properties (i.e., syllabic frequency) constrain orthographic output via a lexical route, at least, in Chinese written production. However, the SF effect over repetitions was divergent in both modalities: it was significant in the former two repetitions in spoken whereas it was significant in the second repetition only in written. Due to the fragility of the SF effect in writing, we suggest that the phonological influence in handwritten production is not mandatory and universal, and it is modulated by experimental manipulations. This provides evidence for the orthographic autonomy hypothesis, rather than the phonological mediation hypothesis. The absence of an interaction between WF and SF showed that the SF effect is independent of the WF effect in spoken and written output modalities. The implications of these results on written production models are discussed.

This paper explored the determinants of picture-naming latency. Using a picture naming task and 5-point scale assessment method, we measured name agreement, familiarity, image agreement, visual complexity and word length of 311 pictures. Correlation analyses indicated that concept agreement, familiarity, image agreement and word frequency were the major determinants of picture-naming latency. Stepwise multiple regressions were carried out on the picture-naming latency. The R square of regression equation was 55.5%. On the basis of the naming latency, the pictures were divided into quintiles. These measures will prove valuable information in future studies of picture naming and other cognitive processes

How is information transmitted across semantic and phonological levels in spoken word production? Recent evidence from speakers of Western languages such as English and Dutch suggests non-discrete transmission, but it is not clear whether this view can be generalized to other languages such as Mandarin, given potential differences in phonological encoding across languages. The present study used Mandarin speakers and combined a behavioral picture-word interference task with event-related potentials. The design factorially crossed semantic and phonological relatedness. Results showed semantic and phonological effects both in behavioral and electrophysiological measurements, with statistical additivity in latencies, and discrete time signatures (250-450 ms and 450-600 ms after picture onset for the semantic and phonological condition, respectively). Overall, results suggest that in Mandarin spoken production, information is transmitted from semantic to phonological levels in a sequential fashion. Hence, temporal signatures associated with spoken word production might differ depending on target language.