Spatio-temporal Patterns Of Cortical Activity Research Articles

Sensory experience selectively reorganizes the late component of evoked responses.

In response to sensory stimulation, the cortex exhibits an early transient response followed by late and slower activation. Recent studies suggest that the early component represents features of the stimulus while the late component is associated with stimulus perception. Although very informative, these studies only focus on the amplitude of the evoked responses to study its relationship with sensory perception. In this work, we expand upon the study of how patterns of evoked and spontaneous activity are modified by experience at the mesoscale level using voltage and extracellular glutamate transient recordings over widespread regions of mouse dorsal neocortex. We find that repeated tactile or auditory stimulation selectively modifies the spatiotemporal patterns of cortical activity, mainly of the late evoked response in anesthetized mice injected with amphetamine and also in awake mice. This modification lasted up to 60min and results in an increase in the amplitude of the late response after repeated stimulation and in an increase in the similarity between the spatiotemporal patterns of the late early evoked response. This similarity increase occurs only for the evoked responses of the sensory modality that received the repeated stimulation. Thus, this selective long-lasting spatiotemporal modification of the cortical activity patterns might provide evidence that evoked responses are a cortex-wide phenomenon. This work opens new questions about how perception-related cortical activity changes with sensory experience across the cortex.

Analysis and Model of Cortical Slow Waves Acquired with Optical Techniques.

Slow waves (SWs) are spatio-temporal patterns of cortical activity that occur both during natural sleep and anesthesia and are preserved across species. Even though electrophysiological recordings have been largely used to characterize brain states, they are limited in the spatial resolution and cannot target specific neuronal population. Recently, large-scale optical imaging techniques coupled with functional indicators overcame these restrictions, and new pipelines of analysis and novel approaches of SWs modelling are needed to extract relevant features of the spatio-temporal dynamics of SWs from these highly spatially resolved data-sets. Here we combined wide-field fluorescence microscopy and a transgenic mouse model expressing a calcium indicator (GCaMP6f) in excitatory neurons to study SW propagation over the meso-scale under ketamine anesthesia. We developed a versatile analysis pipeline to identify and quantify the spatio-temporal propagation of the SWs. Moreover, we designed a computational simulator based on a simple theoretical model, which takes into account the statistics of neuronal activity, the response of fluorescence proteins and the slow waves dynamics. The simulator was capable of synthesizing artificial signals that could reliably reproduce several features of the SWs observed in vivo, thus enabling a calibration tool for the analysis pipeline. Comparison of experimental and simulated data shows the robustness of the analysis tools and its potential to uncover mechanistic insights of the Slow Wave Activity (SWA).

Cortical activation in generalized seizures.

Patients with generalized epilepsy exhibit different epileptiform events including asymptomatic interictal spikes (IS), absence seizures with spike-wave discharges (SWDs), and myoclonic seizures (MS). Our objective was to determine the spatiotemporal patterns of cortical activation in SWDs, IS, and MS in the Gabra1+/A322D juvenile myoclonic epilepsy mouse. We fabricated affordable, flexible high-density electroencephalography (HdEEG) arrays and recorded spontaneous SWD, IS, and MS with video/HdEEG. We determined differences among the events in amplitude spectral density (ASD) in the δ/θ/α/β/γ frequency bands at baseline (3.5-4.0seconds before the first spike time, t0 ) and the prespike period (0.1-0.5seconds before t0 ), and we elucidated the spatiotemporal activation during the t0 spike. All three events had an increase in ASD between baseline and prespike in at least one frequency band. During prespike, MS had the largest δ-band ASD, but SWD had the greatest α/β/γ band ASD. For all three events, the ASD was largest in the anterior regions. The t0 spike voltage was also greatest in the anterior regions for all three events and IS and MS had larger voltages than SWD. From 7.5 to 17.5msec after t0 , MS had greater voltage than IS and SWD, and maximal voltage was in the posterior parietal region. Changes in spectral density from baseline to prespike indicate that none of these generalized events are instantaneous or entirely unpredictable. Prominent engagement of anterior cortical regions during prespike and at t0 suggest that common anterior neural circuits participate in each event. Differences in prespike ASD signify that although the events may engage similar brain regions, they may arise from distinct proictal states with different neuronal activity or connectivity. Prolonged activation of the posterior parietal area in MS suggests that posterior circuits contribute to the myoclonic jerk. Together, these findings identify brain regions and processes that could be specifically targeted for further recording and modulation.

Spatio-temporal characteristics of population responses evoked by microstimulation in the barrel cortex

Intra-cortical microstimulation (ICMS) is a widely used technique to artificially stimulate cortical tissue. This method revealed functional maps and provided causal links between neuronal activity and cognitive, sensory or motor functions. The effects of ICMS on neural activity depend on stimulation parameters. Past studies investigated the effects of stimulation frequency mainly at the behavioral or motor level. Therefore the direct effect of frequency stimulation on the evoked spatio-temporal patterns of cortical activity is largely unknown. To study this question we used voltage-sensitive dye imaging to measure the population response in the barrel cortex of anesthetized rats evoked by high frequency stimulation (HFS), a lower frequency stimulation (LFS) of the same duration or a single pulse stimulation. We found that single pulse and short trains of ICMS induced cortical activity extending over few mm. HFS evoked a lower population response during the sustained response and showed a smaller activation across time and space compared with LFS. Finally the evoked population response started near the electrode site and spread horizontally at a propagation velocity in accordance with horizontal connections. In summary, HFS was less effective in cortical activation compared to LFS although HFS had 5 fold more energy than LFS.

Processing of auditory novelty across the cortical hierarchy: An intracranial electrophysiology study

Under the predictive coding hypothesis, specific spatiotemporal patterns of cortical activation are postulated to occur during sensory processing as expectations generate feedback predictions and prediction errors generate feedforward signals. Establishing experimental evidence for this information flow within cortical hierarchy has been difficult, especially in humans, due to spatial and temporal limitations of non-invasive measures of cortical activity. This study investigated cortical responses to auditory novelty using the local/global deviant paradigm, which engages the hierarchical network underlying auditory predictive coding over short (‘local deviance’; LD) and long (‘global deviance’; GD) time scales. Electrocorticographic responses to auditory stimuli were obtained in neurosurgical patients from regions of interest (ROIs) including auditory, auditory-related and prefrontal cortex. LD and GD effects were assayed in averaged evoked potential (AEP) and high gamma (70–150 Hz) signals, the former likely dominated by local synaptic currents and the latter largely reflecting local spiking activity. AEP LD effects were distributed across all ROIs, with greatest percentage of significant sites in core and non-core auditory cortex. High gamma LD effects were localized primarily to auditory cortex in the superior temporal plane and on the lateral surface of the superior temporal gyrus (STG). LD effects exhibited progressively longer latencies in core, non-core, auditory-related and prefrontal cortices, consistent with feedforward signaling. The spatial distribution of AEP GD effects overlapped that of LD effects, but high gamma GD effects were more restricted to non-core areas. High gamma GD effects had shortest latencies in STG and preceded AEP GD effects in most ROIs. This latency profile, along with the paucity of high gamma GD effects in the superior temporal plane, suggest that the STG plays a prominent role in initiating novelty detection signals over long time scales. Thus, the data demonstrate distinct patterns of information flow in human cortex associated with auditory novelty detection over multiple time scales.

Heightened eating drive and visual food stimuli attenuate central nociceptive processing.

Hunger and pain are basic drives that compete for a behavioral response when experienced together. To investigate the cortical processes underlying hunger-pain interactions, we manipulated participants' hunger and presented photographs of appetizing food or inedible objects in combination with painful laser stimuli. Fourteen healthy participants completed two EEG sessions: one after an overnight fast, the other following a large breakfast. Spatio-temporal patterns of cortical activation underlying the hunger-pain competition were explored with 128-channel EEG recordings and source dipole analysis of laser-evoked potentials (LEPs). We found that initial pain ratings were temporarily reduced when participants were hungry compared with fed. Source activity in parahippocampal gyrus was weaker when participants were hungry, and activations of operculo-insular cortex, anterior cingulate cortex, parahippocampal gyrus, and cerebellum were smaller in the context of appetitive food photographs than in that of inedible object photographs. Cortical processing of noxious stimuli in pain-related brain structures is reduced and pain temporarily attenuated when people are hungry or passively viewing food photographs, suggesting a possible interaction between the opposing motivational forces of the eating drive and pain.

Cortical organization of language pathways in children with non-localized cryptogenic epilepsy.

Children with a history of epilepsy are almost six times more likely than their unaffected siblings to be referred for speech or language therapy. However, the abnormalities in neural pathway that cause these delays are poorly understood. We recorded evoked fields using whole-head magnetoencephalography during real and non-word visual and auditory rhyme tasks in 15 children with non-localized cryptogenic epilepsy. Basic phonological and orthographic language skills were assessed using Woodcock–Johnson Test of Achievement subtests. Dynamic statistical parameter mapping was used with individual participant magnetic resonance images. Significant cortical activity was visualized on average and performance weighted maps. For the auditory rhyme tasks, bilateral primary and secondary auditory cortices, the superior temporal sulcus, and insular cortex were activated early with later increases in left hemisphere activity. Visual rhyme tasks evoked early bilateral primary and secondary occipital cortical and angular gyri activity followed by later activation of the planum temporale and supramarginal gyri and the left ventral occipitotemporal area. For the auditory rhyme tasks, performance weighted maps demonstrated that early right hemisphere activation was associated with poorer reading skills while later activity was associated with better reading skills; for the left hemisphere, greater early activation of the secondary auditory cortex, including the planum temporale, was related to better reading skills while relatively later activation of these areas was associated with poorer reading skills. For the visual rhyme tasks, greater activity in the bilateral ventral occipitotemporal and insular areas and angular and supramarginal gyri were associated with better performance. These data suggest that spatiotemporal cortical activation patterns are associated with variations in language performance in non-localized cryptogenic epilepsy.

Low frequency mu-like activity characterizes cortical rhythms in epilepsy due to ring chromosome 20

ObjectivesTo evaluate the spectral and spatial features of the cortical rhythms in patients affected by ring chromosome 20 – [r(20)]-syndrome. MethodsTwelve patients with [r(20)] syndrome were studied. As controls we enrolled 12 patients with idiopathic generalized epilepsy (IGE) and 12 healthy volunteers (HV). Blind source separation, spectral analyses and source reconstruction were applied in all cases in order to identify reliable spatio-temporal patterns of cortical activity. ResultsA theta–delta EEG rhythm was identified in [r(20)] patients, with spectral peak ranging between 3 and 7Hz and whose generators mapped over the sensory-motor cortices. A second peak laying at a frequency about double with respect to the first one was present in 6 cases. Analogue methodological approach in HV and IGE groups failed to show similar findings. ConclusionsEEG of [r(20)] patients reveals the existence of a highly reproducible EEG pattern arising from the sensory-motor system. SignificanceThe recognition of this peculiar EEG pattern could help the diagnostic work-up. Additionally, our findings supports the existence of a parallelism between this EEG trait and the physiological “mu” rhythm which is generate by the sensory-motor system. Such link suggests a sensory-motor system dysfunction in [r(20)] patients.

Visual Experience Modulates Spatio-Temporal Dynamics of Circuit Activation

Persistent reduction in sensory drive in early development results in multiple plastic changes of different cortical synapses. How these experience-dependent modifications affect the spatio-temporal dynamics of signal propagation in neocortical circuits is poorly understood. Here we demonstrate that brief visual deprivation significantly affects the propagation of electrical signals in the primary visual cortex. The spatio-temporal spread of circuit activation upon direct stimulation of its input layer (Layer 4) is reduced, as is the activation of L2/3 – the main recipient of the output from L4. Our data suggest that the decrease in spatio-temporal activation of L2/3 depends on reduced L4 output, and is not intrinsically generated within L2/3. The data shown here suggest that changes in the synaptic components of the visual cortical circuit result not only in alteration of local integration of excitatory and inhibitory inputs, but also in a significant decrease in overall circuit activation. Furthermore, our data indicate a differential effect of visual deprivation on L4 and L2/3, suggesting that while feedforward activation of L2/3 is reduced, its activation by long range, within layer inputs is unaltered. Thus, brief visual deprivation induces experience-dependent circuit re-organization by modulating not only circuit excitability, but also the spatio-temporal patterns of cortical activation within and between layers.

Comparison of brain mechanisms underlying the processing of Chinese characters and pseudo‐characters: An event‐related potential study

Most Chinese characters are composed of a semantic radical on the left and a phonetic radical on the right. The semantic radical provides the semantic information; the phonetic radical provides information concerning the pronunciation of the whole character. The pseudo‐characters in the study consisted of different sub‐lexical parts of real Chinese characters and consequently they also had the semantic radical and the phonetic radical. But they were not readable and had no actual meaning. In order to investigate the spatiotemporal cortical activation patterns underlying the orthographic, phonological and semantic processing of Chinese characters, we used event‐related brain potentials (ERPs) to explore the processing of Chinese characters and pseudo‐characters when 14 healthy Chinese college students viewed the characters passively. Results showed that both Chinese characters and pseudo‐characters elicited an evident negative potential peaking around 120 ms (N120), which appeared to reflect initial orthographic distinction and evaluation. Then, Chinese pseudo‐characters elicited a more positive ERP deflection (P220) than did Chinese characters 200–250 ms after onset of the stimuli. It was similar to the recognition potential (RP) and might reflect the integration processes of phonological and semantic processing on the basis of early orthographic information. Dipole source analysis of the difference wave (pseudo‐characters minus characters) indicated that a generator localized in the left temporal‐occipital junction contributed to this effect, which was possibly related to phonological and perceptual–semantic information integration. Between 350–450 ms, a greater negativity (N360) in pseudo‐characters as compared to characters was found over midline fronto‐central scalp regions. Dipole analysis localized the generator of N360 in the right parahippocampal cortex. Therefore, the N360 might be an N400 component and reflect the higher‐level semantic activation on the basis of orthographic, phonological and perceptual–semantic processing.

The vase–face illusion seen by the brain: An event-related brain potentials study

In the present study, we provided a cue (face or vase) beforehand to make subjects attend the target stimuli in the vase–face ambiguous figure, and compared the spatiotemporal cortical activation patterns underlying face (face–face) or vase (vase–vase) processing to the ambiguous figure using high-density (64-channel) event-related brain potential (ERP) recordings. Scalp ERP analysis found that the anterior N100, P160, N320 and posterior P100 and N160 were elicited by the face–face and vase–vase responses. The results of the ANOVAs showed that the anterior N100 and N320 elicited by the vase–vase response were more negative than the face–face response. The anterior N100 might reflect deployment of attention (conscious effort) to identify the target stimuli (face or vase) in early processing of the ambiguous figure, and the N320 might be the reversal negativity (RN) and was involved in involuntary perceptual reversals (from face to vase or reverse). Moreover, the mean amplitude (a late positive component: LPC) between 350 and 450 ms of the face–face response was larger than the vase–vase response on positive orientation over the front-central scalp regions. This result might support the view that the LPC reflect post-perceptual processing and indicated that the perceptual reversal of the vase–face illusion is influenced by top-down control.

Spatiotemporal Patterns of Cortical Activity with Bilateral Cochlear Implants in Congenital Deafness

Congenital deafness affects developmental processes in the auditory cortex. In this study, local field potentials (LFPs) were mapped at the cortical surface with microelectrodes in response to cochlear implant stimulation. LFPs were compared between hearing controls and congenitally deaf cats (CDCs). Pulsatile electrical stimulation initially evoked cortical activity in the rostral parts of the primary auditory field (A1). This progressed both in the approximate dorsoventral direction (along the isofrequency stripe) and in the rostrocaudal direction. The dorsal branch of the wavefront split into a caudal branch (propagating in A1) and another smaller one propagating rostrally into the AAF (anterior auditory field). After the front reached the caudal border of A1, a "reflection wave" appeared, propagating back rostrally. In total, the waves took approximately 13-15 ms to propagate along A1 and return back. In CDCs, the propagation pattern was significantly disturbed, with a more synchronous activation of distant cortical regions. The maps obtained from contralateral and ipsilateral stimulation overlapped in both groups of animals. Although controls showed differences in the latency-amplitude patterns, cortical waves evoked by contralateral and ipsilateral stimulation were more similar in CDCs. Additionally, in controls, LFPs with contralateral and ipsilateral stimulation were more similar in caudal A1 than in rostral A1. This dichotomy was lost in deaf animals. In conclusion, propagating cortical waves are specific for the contralateral ear, they are affected by auditory deprivation, and the specificity of the cortex for stimulation of the contralateral ear is reduced by deprivation.

Neural mechanisms underlying the processing of Chinese and English words in a word generation task: An event‐related potential study

Event-related brain potentials (ERPs) were used to examine the spatiotemporal cortical activation patterns underlying Chinese and English word generation (forming a new word by adding a stroke or a letter to an old Chinese or English word) for low- and high-proficiency Chinese-English bilinguals. The results revealed that early visual perceptual processing and word identification were similar between the two languages for the N120 and P220 waveforms. However, a greater negative potential (N250-350) was associated with Chinese words than with English words between 250 and 350 ms. Subsequently, for fluent Chinese bilinguals, Chinese words elicited a more positive ERP deflection (LPC) than did English words between 350 and 800 ms. The differences in ERP components between Chinese and English words indicates that there might be a real difference in the processing demands between these languages, and that the processing of English might be affected by the proficiency of the second language.

Spatiotemporal Signatures of Large-Scale Synfire Chains for Speech Processing as Revealed by MEG

We report a new brain signature of memory trace activation in the human brain revealed by magnetoencephalography and distributed source localization. Spatiotemporal patterns of cortical activation can be picked up in the time course of source images underlying magnetic brain responses to speech and noise stimuli, especially the generators of the magnetic mismatch negativity. We found that acoustic signals perceived as speech elicited a well-defined spatiotemporal pattern of sequential activation of superior–temporal and inferior–frontal cortex, whereas the same identical stimuli, when perceived as noise, did not elicit temporally structured activation. Strength of local sources constituting large-scale spatiotemporal patterns reflected additional lexical and syntactic features of speech. Morphological processing of the critical sound as verb inflection led to particularly pronounced early left inferior–frontal activation, whereas the same sound functioning as inflectional affix of a noun activated superior–temporal cortex more strongly. We conclude that precisely timed spatiotemporal patterns involving specific cortical areas may represent a brain code of memory circuit activation. These spatiotemporal patterns are best explained in terms of synfire mechanisms linking neuronal populations in different cortical areas. The large-scale synfire chains appear to reflect the processing of stimuli together with the context-dependent perceptual and cognitive information bound to them.

Touch or pain? Spatio-temporal patterns of cortical fMRI activity following brief mechanical stimuli

Most imaging studies on the human pain system have concentrated so far on the spatial distribution of pain-related activity. In the present study, we investigated similarities and differences between the spatial and temporal patterns of brain activity related to touch vs. pain perception. To this end, we adopted an event-related functional magnetic resonance imaging (fMRI) paradigm allowing us to separately assess the activity related to stimulus anticipation, perception, and coding. The fMRI signal increases following brief mechanical noxious or non-noxious stimulation of the hand dorsum were largely overlapping in the contralateral and ipsilateral hemispheres, including portions of the parietal, insular, frontal and cingulate cortices. Higher activity following noxious stimulation was found in the contralateral mid-anterior insular cortex, in the anterior mid-cingulate cortex (aMCC) and in the adjacent dorso-medial frontal cortex. Significant decreases in fMRI signals following both tactile and painful stimuli were found in perigenual cingulate (pACC)/medial prefrontal cortex (MPF) and in the posterior cingulate/precuneus/paracentral lobule; more intense decreases were found in the pACC/MPF following painful stimuli. fMRI signal increases in the contralateral insula and in aMCC, but not in the parietal cortex, were more prolonged following painful than tactile stimuli. Moreover, a second peak of signal increases (albeit of lower intensity) was found in anterior insula and aMCC during pain intensity rating. These results show specific spatio-temporal patterns of cortical activity related to processing noxious vs. non-noxious mechanical stimuli.

Objective phonological and subjective perceptual characteristics of syllables modulate spatiotemporal patterns of superior temporal gyrus activity

Natural consonant–vowel syllables are reliably classified by most listeners as voiced or voiceless. However, our previous research [Liederman, J., Frye, R., Fisher, J.M., Greenwood, K., Alexander, R., 2005. A temporally dynamic context effect that disrupts voice onset time discrimination of rapidly successive stimuli. Psychon Bull Rev. 12, 380–386] suggests that among synthetic stimuli varying systematically in voice onset time (VOT), syllables that are classified reliably as voiceless are nonetheless perceived differently within and between listeners. This perceptual ambiguity was measured by variation in the accuracy of matching two identical stimuli presented in rapid succession. In the current experiment, we used magnetoencephalography (MEG) to examine the differential contribution of objective (i.e., VOT) and subjective (i.e., perceptual ambiguity) acoustic features on speech processing. Distributed source models estimated cortical activation within two regions of interest in the superior temporal gyrus (STG) and one in the inferior frontal gyrus. These regions were differentially modulated by VOT and perceptual ambiguity. Ambiguity strongly influenced lateralization of activation; however, the influence on lateralization was different in the anterior and middle/posterior portions of the STG. The influence of ambiguity on the relative amplitude of activity in the right and left anterior STG activity depended on VOT, whereas that of middle/posterior portions of the STG did not. These data support the idea that early cortical responses are bilaterally distributed whereas late processes are lateralized to the dominant hemisphere and support a “how/what” dual-stream auditory model. This study helps to clarify the role of the anterior STG, especially in the right hemisphere, in syllable perception. Moreover, our results demonstrate that both objective phonological and subjective perceptual characteristics of syllables independently modulate spatiotemporal patterns of cortical activation.

Brain mechanisms underlying visual perception and visual mental imagery of Chinese pseudo-characters: An event-related potential study

The aim of this study was to compare spatiotemporal cortical activation patterns that underlie the processing of visual perception and visual mental imagery of Chinese pseudo-characters. Event-related brain potentials (ERPs) were measured when 14 healthy Chinese college students performed a pseudo-character stroke judgment task. Results showed that visual mental imagery elicited a more negative ERP deflection (N240) than visual perception in the 150- to 250-ms time window after onset of the stimuli. Maps of the difference wave (mental imagery-visual perception) showed strong activity in the right frontal regions. Dipole analysis revealed that the generator of N240 was localized in the right parahippocampal cortex and possibly related to forming a visual mental imagery of the pseudo-character on the basis of initial stimuli identification and classification. Then in the time window between 400 and 600 ms, a greater negativity (N520) in visual mental imagery as compared to visual perception was detected over the right frontocentral scalp regions. Moreover, dipole source analysis of the difference wave (mental imagery-visual perception) indicated that a generator was localized in the right temporal-occipital junction (BA18), which appeared to reflect the cognitive processes of character reconstruction and stroke searching in the visual mental imagery of the pseudo-character. The results showed that the visual mental imagery of Chinese pseudo-characters might be related to the right parahippocampal cortex, and activation of the right temporal-occipital junction was possibly related to accomplishing the stroke judgment task in the visual mental imagery.

Propagating Waves in Visual Cortex

Spatiotemporal patterns of cortical activation occur superimposed upon sensory, motor, and cognitive maps. The papers by Benucci et al. and Xu et al. in this issue of Neuron demonstrate that visual responses propagate in space and may serve to link different visual areas. This is an important step toward understanding how cortical maps relate to activation patterns, a prerequisite to understanding visual cortex function.

Comparison of spatiotemporal cortical activation pattern during visual perception of Korean, English, Chinese words: An event-related potential study

The aim of this study was to compare spatiotemporal cortical activation patterns during the visual perception of Korean, English, and Chinese words. The comparison of these three languages offers an opportunity to study the effect of written forms on cortical processing of visually presented words, because of partial similarity/difference among words of these languages, and the familiarity of native Koreans with these three languages at the word level. Single-character words and pictograms were excluded from the stimuli in order to activate neuronal circuitries that are involved only in word perception. Since a variety of cerebral processes are sequentially evoked during visual word perception, a high-temporal resolution is required and thus we utilized event-related potential (ERP) obtained from high-density electroencephalograms. The differences and similarities observed from statistical analyses of ERP amplitudes, the correlation between ERP amplitudes and response times, and the patterns of current source density, appear to be in line with demands of visual and semantic analysis resulting from the characteristics of each language, and the expected task difficulties for native Korean subjects.

Neuromagnetic brain responses to words from semantic sub- and supercategories

BackgroundWe explored spatio-temporal patterns of cortical activity evoked by written words from super-ordinate and sub-ordinate semantic categories and hoped to find a differential cortical and/or temporal distribution of the brain response depending on the level of the categories. Twenty-three subjects saw 360 words belonging to six sub-ordinate categories (mammals, birds, fish, fruit, flowers, trees) within two super-ordinate categories (fauna, flora). Visually evoked magnetic fields were determined from whole-head (148-sensor) magnetoencephalography and analyzed in the source space (Minimum Norm Estimate).ResultsActivity (MNE amplitudes) 100–150 ms after stimulus onset in the left occipito-temporal area distinguished super-ordinate categories, while later activity (300–550 ms) in the left temporal area distinguished the six sub-ordinate categories.ConclusionOur results document temporally and spatially distinct processing and representation of words according to their categorical information. If further studies can rule out possible confounds then our results may help constructing a theory about the internal structure of entries in the mental lexicon and its access.