Brain activation for lexical decision and reading aloud: two sides of the same coin?

This functional magnetic resonance imaging study compared the neuronal implementation of word and pseudoword processing during two commonly used word recognition tasks: lexical decision and reading aloud. In the lexical decision task, participants made a finger-press response to indicate whether a visually presented letter string is a word or a pseudoword (e.g., "paple"). In the reading-aloud task, participants read aloud visually presented words and pseudowords. The same sets of words and pseudowords were used for both tasks. This enabled us to look for the effects of task (lexical decision vs. reading aloud), lexicality (words vs. nonwords), and the interaction of lexicality with task. We found very similar patterns of activation for lexical decision and reading aloud in areas associated with word recognition and lexical retrieval (e.g., left fusiform gyrus, posterior temporal cortex, pars opercularis, and bilateral insulae), but task differences were observed bilaterally in sensorimotor areas. Lexical decision increased activation in areas associated with decision making and finger tapping (bilateral postcentral gyri, supplementary motor area, and right cerebellum), whereas reading aloud increased activation in areas associated with articulation and hearing the sound of the spoken response (bilateral precentral gyri, superior temporal gyri, and posterior cerebellum). The effect of lexicality (pseudoword vs. words) was also remarkably consistent across tasks. Nevertheless, increased activation for pseudowords relative to words was greater in the left precentral cortex for reading than lexical decision, and greater in the right inferior frontal cortex for lexical decision than reading. We attribute these effects to differences in the demands on speech production and decision-making processes, respectively.     

Spectral-temporal analysis of cortical oscillations during lexical processing

We investigated the oscillatory neural correlates of auditory lexical processing in healthy adults. Synthetic aperture magnetometry was used to characterize the timing of event-related desynchronization (ERD)/event-related synchronization (ERS) in superior temporal gyri following low-frequency and high-frequency words in contrast to nonwords. ERS and ERD responses were found with both word and nonword stimuli. Analysis of power revealed significantly elevated θ-α range (6-14 Hz) ERD in response to words compared with nonwords (left hemisphere: 390-945 ms poststimulus). Furthermore, a burst of ERS in the γ band (40-50 Hz, centered at 410 ms poststimulus) distinguished high-frequency and low-frequency words, and also displayed left-hemispheric enhancement following words. Results demonstrate a clear neural correlate of lexical access and provide a basis for further study of spectral-temporal brain activity during language processing.     

Lists with and without Syntax: A New Approach to Measuring the Neural Processing of Syntax

In the neurobiology of syntax, a methodological challenge is to vary syntax while holding semantics constant. Changes in syntactic structure usually correlate with changes in meaning. We approached this challenge from a new angle. We deployed word lists—typically, the unstructured control in studies of syntax—as both test and control stimuli. Three-noun lists (“lamps, dolls, guitars”) were embedded in sentences (“The eccentric man hoarded lamps, dolls, guitars…”) and in longer lists (“forks, pen, toilet, rodeo, lamps, dolls, guitars…”). This allowed us to minimize contributions from lexical semantics and local phrasal combinatorics: the same words occurred in both conditions, and in neither case did the list items locally compose into phrases (e.g., “lamps” and “dolls” do not form a phrase). Crucially, the list partakes in a syntactic tree in one case but not the other. Lists-in-sentences increased source-localized MEG activity at ∼250–300 ms from each of the list item onsets in the left inferior frontal cortex, at ∼300–350 ms in the left anterior temporal lobe and, most reliably, at ∼330–400 ms in left posterior temporal cortex. In contrast, the main effects of semantic association strength, which we also varied, localized in the left temporoparietal cortex, with high associations increasing activity at ∼400 ms. This dissociation offers a novel characterization of the structure versus word meaning contrast in the brain: the frontotemporal network that is familiar from studies of sentence processing can be driven by the sheer presence of global sentence structure, while associative semantics has a more posterior neural signature.SIGNIFICANCE STATEMENT Human languages all have a syntax, which both enables the infinitude of linguistic creativity and determines what is grammatical in a language. The neurobiology of syntactic processing has, however, been challenging to characterize despite decades of study. One reason is pure manipulations of syntax are difficult to design. The approach here offers a novel control of two variables that are notoriously hard to keep constant when syntax is manipulated: word meaning and phrasal combinatorics. The same noun lists occurred inside longer lists and sentences, while semantic associations also varied. Our MEG results show that classic frontotemporal language regions can be driven by sentence structure even when local semantic contributions are absent. In contrast, the left temporoparietal junction tracks associative relationships.     

Pseudohomophone effects provide evidence of early lexico-phonological processing in visual word recognition

Previous research using event-related brain potentials (ERPs) suggested that phonological processing in visual word recognition occurs rather late, typically after semantic or syntactic processing. Here, we show that phonological activation in visual word recognition can be observed much earlier. Using a lexical decision task, we show that ERPs to pseudohomophones (PsHs) (e.g., ROZE) differed from well-matched spelling controls (e.g., ROFE) as early as 150 ms (P150) after stimulus onset. The PsH effect occurred as early as the word frequency effect suggesting that phonological activation occurs early enough to influence lexical access. Low-resolution electromagnetic tomography analysis (LORETA) revealed that left temporoparietal and right frontotemporal areas are the likely brain regions associated with the processing of phonological information at the lexical level. Altogether, the results show that phonological processes are activated early in visual word recognition and play an important role in lexical access.     

Online processing of moral transgressions: ERP evidence for spontaneous evaluation

Experimental studies using fictional moral dilemmas indicate that both automatic emotional processes and controlled cognitive processes contribute to moral judgments. However, not much is known about how people process socio-normative violations that are more common to their everyday life nor the time-course of these processes. Thus, we recorded participants’ electrical brain activity while they were reading vignettes that either contained morally acceptable vs unacceptable information or text materials that contained information which was either consistent or inconsistent with their general world knowledge. A first event-related brain potential (ERP) positivity peaking at ∼200 ms after critical word onset (P200) was larger when this word involved a socio-normative or knowledge-based violation. Subsequently, knowledge-inconsistent words triggered a larger centroparietal ERP negativity at ∼320 ms (N400), indicating an influence on meaning construction. In contrast, a larger ERP positivity (larger late positivity), which also started at ∼320 ms after critical word onset, was elicited by morally unacceptable compared with acceptable words. We take this ERP positivity to reflect an implicit evaluative (good–bad) categorization process that is engaged during the online processing of moral transgressions.     

The signer and the sign: Cortical correlates of person identity and language processing from point-light displays

In this study, the first to explore the cortical correlates of signed language (SL) processing under point-light display conditions, the observer identified either a signer or a lexical sign from a display in which different signers were seen producing a number of different individual signs. Many of the regions activated by point-light under these conditions replicated those previously reported for full-image displays, including regions within the inferior temporal cortex that are specialised for face and body-part identification, although such body parts were invisible in the display. Right frontal regions were also recruited – a pattern not usually seen in full-image SL processing. This activation may reflect the recruitment of information about person identity from the reduced display. A direct comparison of identify-signer and identify-sign conditions showed these tasks relied to a different extent on the posterior inferior regions. Signer identification elicited greater activation than sign identification in (bilateral) inferior temporal gyri (BA 37/19), fusiform gyri (BA 37), middle and posterior portions of the middle temporal gyri (BAs 37 and 19), and superior temporal gyri (BA 22 and 42). Right inferior frontal cortex was a further focus of differential activation (signer > sign).These findings suggest that the neural systems supporting point-light displays for the processing of SL rely on a cortical network including areas of the inferior temporal cortex specialized for face and body identification. While this might be predicted from other studies of whole body point-light actions (Vaina, Solomon, Chowdhury, Sinha, & Belliveau, 2001) it is not predicted from the perspective of spoken language processing, where voice characteristics and speech content recruit distinct cortical regions (Stevens, 2004) in addition to a common network. In this respect, our findings contrast with studies of voice/speech recognition (Von Kriegstein, Kleinschmidt, Sterzer, & Giraud, 2005). Inferior temporal regions associated with the visual recognition of a person appear to be required during SL processing, for both carrier and content information.     

Lexical and syntactic representations in the brain: an fMRI investigation with multi-voxel pattern analyses

Work in theoretical linguistics and psycholinguistics suggests that human linguistic knowledge forms a continuum between individual lexical items and abstract syntactic representations, with most linguistic representations falling between the two extremes and taking the form of lexical items stored together with the syntactic/semantic contexts in which they frequently occur. Neuroimaging evidence further suggests that no brain region is selectively sensitive to only lexical information or only syntactic information. Instead, all the key brain regions that support high-level linguistic processing have been implicated in both lexical and syntactic processing, suggesting that our linguistic knowledge is plausibly represented in a distributed fashion in these brain regions. Given this distributed nature of linguistic representations, multi-voxel pattern analyses (MVPAs) can help uncover important functional properties of the language system. In the current study we use MVPAs to ask two questions: (1) Do language brain regions differ in how robustly they represent lexical vs. syntactic information? and (2) Do any of the language bran regions distinguish between “pure” lexical information (lists of words) and “pure” abstract syntactic information (jabberwocky sentences) in the pattern of activity? We show that lexical information is represented more robustly than syntactic information across many language regions (with no language region showing the opposite pattern), as evidenced by a better discrimination between conditions that differ along the lexical dimension (sentences vs. jabberwocky, and word lists vs. nonword lists) than between conditions that differ along the syntactic dimension (sentences vs. word lists, and jabberwocky vs. nonword lists). This result suggests that lexical information may play a more critical role than syntax in the representation of linguistic meaning. We also show that several language regions reliably discriminate between “pure” lexical information and “pure” abstract syntactic information in their patterns of neural activity.     

Operant conditioning of single unit activity in parietal cortex

Two rhesus monkeys were trained to control firing patterns of single neurons in parietal cortex (areas 1, 2, 3, 5, 7) using an operant task previously applied to the study of precental units. Twenty-four of 56 (43%) postcentral cells were controlled in contrast to 71 of 136 (52%) precentral units from these and 4 other rhesus monkeys. In addition, monkeys were able to drive precentral units to more sustained tonic firing rates than they could parietal units.An analysis of interspike interval (ISI) distributions showed that, in contrast to precentral units with modal ISIs of 25–50 ms, 50% of parietal units have modal ISIs of 2 ms. Such short ISIs may account for fewer postcentral units reaching control criteria for this particular operant task. Other factors that may contribute to the reduced control of postcentral cells are discussed, particularly the more complex afferent connections to parietal units when compared to precentral pyramidal tract neurons.The data indirectly support conclusions from previous studies that imply that operant control of cortical units is peripherally mediated and does not primarily involve a ‘central’ or ‘open loop’ system.     

Cortical gamma‐oscillations modulated by auditory–motor tasks‐intracranial recording in patients with epilepsy

Human activities often involve hand‐motor responses following external auditory–verbal commands. It has been believed that hand movements are predominantly driven by the contralateral primary sensorimotor cortex, whereas auditory–verbal information is processed in both superior temporal gyri. It remains unknown whether cortical activation in the superior temporal gyrus during an auditory–motor task is affected by laterality of hand‐motor responses. Here, event‐related γ‐oscillations were intracranially recorded as quantitative measures of cortical activation; we determined how cortical structures were activated by auditory‐cued movement using each hand in 15 patients with focal epilepsy. Auditory–verbal stimuli elicited augmentation of γ‐oscillations in a posterior portion of the superior temporal gyrus, whereas hand‐motor responses elicited γ‐augmentation in the pre‐ and postcentral gyri. The magnitudes of such γ‐augmentation in the superior temporal, precentral, and postcentral gyri were significantly larger when the hand contralateral to the recorded hemisphere was required to be used for motor responses, compared with when the ipsilateral hand was. The superior temporal gyrus in each hemisphere might play a greater pivotal role when the contralateral hand needs to be used for motor responses, compared with when the ipsilateral hand does. Hum Brain Mapp, 2010. © 2010 Wiley‐Liss, Inc.     

Magnetic resonance and positron emission tomography changes during the clinical progression of Rasmussen encephalitis.

The authors describe serial positron emission tomography (PET) and magnetic resonance imaging (MRI) studies in a patient with pathologically confirmed Rasmussen Encephalitis (RE). Results of initial PET and MRI studies were normal. Subsequent studies showed involvement of the percentral and postcentral gyri and the putamen on PET, and the precentral and postcentral gyri on MRI. Coregistration of PET and MR images showed good correlation between the precentral and postcentral gyri involvement. However, subcortical involvement occurred earlier on PET than on MRI. The authors demonstrate the evolution of changes on PET and MR images in a patient with RE. Despite early pathologic confirmation of RE, there were no definite structural or functional imaging changes on PET or MRI until 3 years after symptom onset. These findings demonstrate the variability of imaging changes in RE, and the need to carefully correlate electro-physiologic and clinical findings to confirm the diagnosis of RE.     

Somatosensory evoked potentials recorded directly from human thalamus and Sm I cortical area.

Somatosensory evoked potentials (SEPs) to median nerve stimulation were recorded from the nucleus ventralis caudalis. They consisted of monophasic or diphasic potentials with mean onset latency of 13.8 ms. More complex SEPs to median nerve stimulation were obtained from the cortex. The SEPs consisted of two major positive waves, P1 and P2, and were recorded over both the precentral and postcentral gyri, suggesting that somatosensory information converges to the motor cortex, probably to be used for the integration of critical motor activity. In two patients, it was noted that the motor representation of facial movements was larger than the correspondent sensory representation on the postcentral gyrus. This larger motor representation of the face and more specifically of the lips and tongue may be related to human acquisition of mimicry and articulation of language.     

Neural correlates of semantic priming for ambiguous words: an event-related fMRI study

We investigated the neural correlates of semantic priming by using event-related fMRI to record blood oxygen level dependent (BOLD) responses while participants performed speeded lexical decisions (word/nonword) on visually presented related versus unrelated prime-target pairs. A long stimulus onset asynchrony of 1000 ms was employed, which allowed for increased controlled processing and selective frequency-based ambiguity priming. Conditions included an ambiguous word prime (e.g. bank) and a target related to its dominant (e.g. money) or subordinate meaning (e.g. river). Compared to an unrelated condition, primed dominant targets were associated with increased activity in the LIFG, the right anterior cingulate and superior temporal gyrus, suggesting postlexical semantic integrative mechanisms, while increased right supramarginal activity for the unrelated condition was consistent with expectancy based priming. Subordinate targets were not primed and were associated with reduced activity primarily in occipitotemporal regions associated with word recognition, which may be consistent with frequency-based meaning suppression. These findings provide new insights into the neural substrates of semantic priming and the functional-anatomic correlates of lexical ambiguity suppression mechanisms.     

Reading fluent speech from talking faces: typical brain networks and individual differences

Listeners are able to extract important linguistic information by viewing the talker's face-a process known as 'speechreading.' Previous studies of speechreading present small closed sets of simple words and their results indicate that visual speech processing engages a wide network of brain regions in the temporal, frontal, and parietal lobes that are likely to underlie multiple stages of the receptive language system. The present study further explored this network in a large group of subjects by presenting naturally spoken sentences which tap the richer complexities of visual speech processing. Four different baselines (blank screen, static face, nonlinguistic facial gurning, and auditory speech) enabled us to determine the hierarchy of neural processing involved in speechreading and to test the claim that visual input reliably accesses sound-based representations in the auditory cortex. In contrast to passively viewing a blank screen, the static-face condition evoked activation bilaterally across the border of the fusiform gyrus and cerebellum, and in the medial superior frontal gyrus and left precentral gyrus (p < .05, whole brain corrected). With the static face as baseline, the gurning face evoked bilateral activation in the motion-sensitive region of the occipital cortex, whereas visual speech additionally engaged the middle temporal gyrus, inferior and middle frontal gyri, and the inferior parietal lobe, particularly in the left hemisphere. These latter regions are implicated in lexical stages of spoken language processing. Although auditory speech generated extensive bilateral activation across both superior and middle temporal gyri, the group-averaged pattern of speechreading activation failed to include any auditory regions along the superior temporal gyrus, suggesting that f luent visual speech does not always involve sound-based coding of the visual input. An important finding from the individual subject analyses was that activation in the superior temporal gyrus did reach significance (p < .001, small-volume corrected) for a subset of the group. Moreover, the extent of the left-sided superior temporal gyrus activity was strongly correlated with speechreading performance. Skilled speechreading was also associated with activations and deactivations in other brain regions, suggesting that individual differences ref lect the efficiency of a circuit linking sensory, perceptual, memory, cognitive, and linguistic processes rather than the operation of a single component process.     

Age‐related microstructural changes in subcortical white matter during postadolescent periods in men revealed by diffusion‐weighted MR imaging

Continuous maturation of cerebral white matter (WM) in the postadolescent period is not fully understood. To elucidate the time course and location of possible postadolescent maturational changes in cerebral WM, we studied 60 healthy male subjects who were in their second to seventh decade using diffusion‐weighted imaging. Mean diffusivity (MD) in subcortical WM was measured in 78 cortical regions in each subject's brain using an automated method. Regression analysis was used to model the age‐related change in MD by either a linear or a quadratic function in each region. Age‐related changes in subcortical MD were best modeled by either a linear function or a quadratic function in 27 regions including language‐related regions, visual or multimodal areas in the bilateral occipital and temporal lobes, limbic areas including the bilateral parahippocampal gyri, and the bilateral postcentral and left precentral gyri. In these regions, the MD rapidly decreased until middle age and thereafter reached a plateau. Our results revealed microstructural changes in local subcortical WM and suggests a continuing maturational process in postoadolescent periods. Hum Brain Mapp 2009. © 2009 Wiley‐Liss, Inc.     

A multicentre study of motor functional connectivity changes in patients with multiple sclerosis

In this multicentre study involving eight European centres, we characterized the spatial pattern of functional connectivity (FC) in the sensorimotor network from 61 right-handed patients with multiple sclerosis (MS) and 74 age-matched healthy subjects assessed with the use of functional magnetic resonance imaging (fMRI) and a simple motor task of their right dominant hand. FC was investigated by using: (i) voxel-wise correlations between the left sensorimotor cortex (SMC) and any other area in the brain; and (ii) bivariate correlations between time series extracted from several regions of interest (ROIs) belonging to the sensorimotor network. Both healthy controls and MS patients had significant FC between the left SMC and several areas of the sensorimotor network, including the bilateral postcentral and precentral gyri, supplementary motor area, middle frontal gyri, insulae, secondary somatosensory cortices, thalami, and right cerebellum. Voxel-wise assessment of FC revealed increased connectivity between the left SMC and the right precentral gyrus, right middle frontal gyrus (MFG) and bilateral postcentral gyri in MS patients as compared with controls. ROI analysis also showed a widespread pattern of altered connectivity, characterized by increased FC between the right MFG, the left insula and the right inferior frontal gyrus in comparison with many regions of the sensorimotor network. These results provide further evidence for increased bihemispheric contributions to motor control in patients with MS relative to healthy controls. They further suggest that multicentre fMRI studies of FC changes are possible, and provide a potential imaging biomarker for use in experimental therapeutic studies directed at enhancing adaptive plasticity in the disease.     

Widespread cortical thinning in children with frontal lobe epilepsy

Purpose: Spread of seizure activity outside the frontal lobe due to cortico‐cortical connections can result in alteration in the cortex beyond the frontal lobe in children with intractable frontal lobe epilepsy (FLE). The aim of this study was to identify regions of reduced cortical thickness in children with intractable FLE. Methods: High‐resolution volumetric T₁‐weighted imaging was performed on 17 children with FLE, who were being evaluated for epilepsy surgery, and 26 age‐matched healthy controls. The cortical thickness of 12 patients with left FLE and 5 patients with right FLE was compared to controls. The clusters of cortical thinning were regressed against age of seizure onset, duration of epilepsy, seizure frequency, and number of medications. Key Findings: In children with left FLE, cortical thinning was present in the left superior frontal, paracentral, precuneus, cingulate, inferior parietal, supramarginal, postcentral, and superior temporal gyri, as well as in the right superior and middle frontal, medial orbitofrontal, supramarginal, postcentral, banks of superior temporal sulcus, and parahippocampal gyri. In children with right FLE, cortical thinning was present in the right precentral, postcentral, transverse temporal, parahippocampal, lingual, and lateral occipital gyri, as well as in the left superior frontal, inferior parietal, postcentral, superior temporal, posterior cingulate, and lingual gyri. In children with left FLE, following exclusion of one outlier, there was no significant association between age at seizure onset, duration of epilepsy, seizure frequency and number of medications with clusters of cortical thinning. In children with right FLE, age at seizure onset, duration of epilepsy, frequency of seizures, and number of medications were not associated with clusters of cortical thinning within the right and left hemispheres. Significance: Cortical changes were present in the frontal and extrafrontal cortex in children with intractable FLE. These changes may be related to spread of seizure activity, large epileptogenic zones involving both frontal and extrafrontal lobes, and development of secondary epileptogenic zones that over time lead to cortical abnormality. Further studies correlating cortical changes with neurocognitive measures are needed to determine if the cortical changes relate to cognitive function.     

Modality independence of word comprehension

Functional magnetic resonance imaging (fMRI) was used to examine the functional anatomy of word comprehension in the auditory and visual modalities of presentation. We asked our subjects to determine if word pairs were semantically associated (e.g., table, chair) and compared this to a reference task where they were asked to judge whether word pairs rhymed (e.g., bank, tank). This comparison showed task-specific and modality-independent activation for semantic processing in the heteromodal cortices of the left inferior frontal gyrus (BA 46, 47) and left middle temporal gyrus (BA 21). There were also modality-specific activations in the fusiform gyrus (BA 37) for written words and in the superior temporal gyrus (BA 22) for spoken words. Our findings are consistent with the hypothesis that word form recognition (lexical encoding) occurs in unimodal cortices and that heteromodal brain regions in the anterior as well as posterior components of the language network subserve word comprehension (semantic decoding).     

Cortical activation during word reading and picture naming in dyslexic and non-reading-impaired children.

OBJECTIVE: In a recent study on picture naming and word reading in dyslexics and control children we found a combination of normal picture retrieval times and severe reading impairments in dyslexics. Therefore, we hypothesize that brain response patterns differ between patients and controls during word reading, but are similar in picture naming as a non-letter mediated task. METHODS: Time course of brain activation was investigated by magnetoencephalography during word reading and picture naming in 9 dyslexic children and 13 age-matched controls (aged 9-10 years). RESULTS: We found 5 consecutive activations spreading from occipito-parietal to temporo-frontal sites. Group differences occurred only during reading: a delayed response in temporal superior and angular gyri at 235-285 ms and absence of activation in anterior temporal and inferior frontal regions at 430-530 ms for dyslexics. CONCLUSIONS: Problems in phonological processing are reflected in delay of early activity and absence of late activity in language related brain regions. From the lack of group differences during picture naming, we conclude the presence of two pathways: a phonological/orthographic one for word reading, which is disturbed in dyslexics, and a visual one for picture naming, which can be unaffected in dyslexics. SIGNIFICANCE: Evidence is provided for different pathways for the processing of letter-mediated and visual-eidetic information. This knowledge may be important for dyslexics in the context of coping with everyday demands and for training of relevant skills.     

Cortical processing of tactile language in a postlingually deaf-blind subject

We compared neural activation detected by magnetoencephalography (MEG) during tactile presentation of words and non-words in a postlingually deaf-blind subject and six normal volunteers. The left postcentral gyrus, bilateral inferior frontal gyri, left posterior temporal lobe, right anterior temporal lobe, bilateral middle occipital gyri were activated when tactile words were presented to the right hand of the deaf-blind subject. This set of activated regions was not observed in the normal volunteers, although activation of several combinations of these regions was detected. Positron emission tomography confirmed the location of the MEG-activated areas in the deaf-blind subject. Our results demonstrated that the deaf-blind subject is heavily involved in interpreting tactile language by enhancing cortical activation of cognitive and semantic processing.