Graph theoretical analysis reveals the functional role of the left ventral occipito-temporal cortex in speech processing.

The left ventral occipito-temporal cortex (left-vOT) plays a key role in reading. Interestingly, the area also responds to speech input, suggesting that it may have other functions beyond written word recognition. Here, we adopt graph theoretical analysis to investigate the left-vOT's functional role in the whole-brain network while participants process spoken sentences in different contexts. Overall, different connectivity measures indicate that the left-vOT acts as an interface enabling the communication between distributed brain regions and sub-networks. During simple speech perception, the left-vOT is systematically part of the visual network and contributes to the communication between neighboring areas, remote areas, and sub-networks, by acting as a local bridge, a global bridge, and a connector, respectively. However, when speech comprehension is explicitly required, the specific functional role of the area and the sub-network to which the left-vOT belongs change and vary with the quality of speech signal and task difficulty. These connectivity patterns provide insightful information on the contribution of the left-vOT in various contexts of language processing beyond its role in reading. They advance our general understanding of the neural mechanisms underlying the flexibility of the language network that adjusts itself according to the processing context.

Inter-individual variability in dorsal stream dynamics during word production.

The current standard model of language production involves a sensorimotor dorsal stream connecting areas in the temporo-parietal junction with those in the inferior frontal gyrus and lateral premotor cortex. These regions have been linked to various aspects of word production such as phonological processing or articulatory programming, primarily through neuropsychological and functional imaging group studies. Most if not all the theoretical descriptions of this model imply that the same network should be identifiable across individual speakers. We tested this hypothesis by quantifying the variability of activation observed across individuals within each dorsal stream anatomical region. This estimate was based on electrical activity recorded directly from the cerebral cortex with millisecond accuracy in awake epileptic patients clinically implanted with intracerebral depth electrodes for pre-surgical diagnosis. Each region's activity was quantified using two different metrics-intra-cerebral evoked related potentials and high gamma activity-at the level of the group, the individual and the recording contact. The two metrics show simultaneous activation of parietal and frontal regions during a picture naming task, in line with models that posit interactive processing during word retrieval. They also reveal different levels of between-patient variability across brain regions, except in core auditory and motor regions. The independence and non-uniformity of cortical activity estimated through the two metrics push the current model towards sub-second and sub-region explorations focused on individualized language speech production. Several hypotheses are considered for this within-region heterogeneity.

Advances in human intracranial electroencephalography research, guidelines and good practices.

Since the second-half of the twentieth century, intracranial electroencephalography (iEEG), including both electrocorticography (ECoG) and stereo-electroencephalography (sEEG), has provided an intimate view into the human brain. At the interface between fundamental research and the clinic, iEEG provides both high temporal resolution and high spatial specificity but comes with constraints, such as the individual's tailored sparsity of electrode sampling. Over the years, researchers in neuroscience developed their practices to make the most of the iEEG approach. Here we offer a critical review of iEEG research practices in a didactic framework for newcomers, as well addressing issues encountered by proficient researchers. The scope is threefold: (i) review common practices in iEEG research, (ii) suggest potential guidelines for working with iEEG data and answer frequently asked questions based on the most widespread practices, and (iii) based on current neurophysiological knowledge and methodologies, pave the way to good practice standards in iEEG research. The organization of this paper follows the steps of iEEG data processing. The first section contextualizes iEEG data collection. The second section focuses on localization of intracranial electrodes. The third section highlights the main pre-processing steps. The fourth section presents iEEG signal analysis methods. The fifth section discusses statistical approaches. The sixth section draws some unique perspectives on iEEG research. Finally, to ensure a consistent nomenclature throughout the manuscript and to align with other guidelines, e.g., Brain Imaging Data Structure (BIDS) and the OHBM Committee on Best Practices in Data Analysis and Sharing (COBIDAS), we provide a glossary to disambiguate terms related to iEEG research.

Motor resonance during linguistic processing as shown by EEG in a naturalistic VR environment.

Embodied cognition studies have shown motor resonance during action language processing, indicating that linguistic representations are at least partially multimodal. However, constraints of this activation linked to linguistic and extra-linguistic context, function and timing have not yet been fully explored. Importantly, embodied cognition binds social and physical contexts to cognition, suggesting that more ecologically valid contexts will yield more valid measures of cognitive processing. Herein, we measured cortical motor activation during language processing in a fully immersive Cave automatic virtual environment (CAVE). EEG was recorded while participants engaged in a Go/No-Go task. They heard action verbs and, for Go trials, performed a corresponding action on a virtual object. ERSP (event-related spectral perturbation) was calculated during verb processing, corresponding to the pattern of power suppression (event-related desynchronization - ERD) and enhancement (event-related synchronization - ERS) relative to the reference interval. Significant ERD emerged during verb processing in both the µ (8-13 Hz) and beta band (20-30 Hz) for both Go and No-Go trials. µ ERD emerged in the 400-500 msec time window, associated with lexical-semantic processing. Greater µ ERD emerged for Go compared to No-Go trials. The present results provide compelling evidence in a naturalistic setting of how motor and linguistic processes interact.

Cortical Dynamics of Semantic Priming and Interference during Word Production: An Intracerebral Study.

Language production requires that semantic representations are mapped to lexical representations on the basis of the ongoing context to select the appropriate words. This mapping is thought to generate two opposing phenomena, "semantic priming," where multiple word candidates are activated, and "interference," where these word activities are differentiated to make a goal-relevant selection. In previous neuroimaging and neurophysiological research, priming and interference have been associated to activity in regions of a left frontotemporal network. Most of such studies relied on recordings that either have high temporal or high spatial resolution, but not both. Here, we employed intracerebral EEG techniques to explore with both high resolutions, the neural activity associated with these phenomena. The data came from nine epileptic patients who were stereotactically implanted for presurgical diagnostics. They performed a cyclic picture-naming task contrasting semantically homogeneous and heterogeneous contexts. Of the 84 brain regions sampled, 39 showed task-evoked activity that was significant and consistent across two patients or more. In nine of these regions, activity was significantly modulated by the semantic manipulation. It was reduced for semantically homogeneous contexts (i.e., priming) in eight of these regions, located in the temporal ventral pathway as well as frontal areas. Conversely, it was increased only in the pre-SMA, notably at an early poststimulus temporal window (200-300 msec) and a preresponse temporal window (700-800 msec). These temporal effects respectively suggest the pre-SMA's role in initial conflict detection (e.g., increased response caution) and in preresponse control. Such roles of the pre-SMA are traditional from a history of neural evidence in simple perceptual tasks, yet are also consistent with recent cognitive lexicosemantic theories that highlight top-down processes in language production. Finally, although no significant semantic modulation was found in the ACC, future intracerebral EEG work should continue to inspect ACC with the pre-SMA.

Grammatical class modulates the (left) inferior frontal gyrus within 100 milliseconds when syntactic context is predictive.

The current study set out to examine the spatiotemporal dynamics of predictive processing during syntactic processing. To do so, we conducted an MEG experiment in which we contrasted MRI-constrained sources elicited by nouns and verbs when they were preceded by a predictive syntactic context (i.e., possessive pronouns for nouns, and personal pronouns for verbs) versus a non-predictive syntactic context (visually matched symbols). The results showed rapid (from ~80 ms onwards) noun-verb differences in the left and (to a lesser extent) right inferior frontal gyri (IFG), but only when those nouns and verbs were preceded by the syntactically predictive context (i.e. their corresponding pronoun). Furthermore, the contrast between possessive and personal pronouns that preceded the rapid noun-verb modulations in the (L)IFG also produced differences in source activation in various regions of the prefrontal cortex (the superior frontal and orbitofrontal cortex). We suggest the data show that syntactic unification manifests very early on during processing in the LIFG. The speed of such syntactic unification operations is hypothesized to be driven by predictive top-down activations stemming from a domain-general network in the prefrontal cortex.

Simultaneous SEEG-MEG-EEG recordings Overcome the SEEG limited spatial sampling.

During presurgical evaluation of pharmacoresistant partial epilepsies, stereoelectroencephalography (SEEG) records interictal and ictal activities directly but is inherently limited in spatial sampling. In contrast, scalp-EEG and MEG are less sensitive but provide a global view on brain activity. Therefore, recording simultaneously these three modalities should provide a better understanding of the underlying brain sources by taking advantage of the different sensitivities of the three recording techniques. We performed trimodal EEG-MEG-SEEG recordings in a 19-year-old woman with pharmacoresistant cryptogenic posterior cortex epilepsy. Sub-continuous and highly focal spikes that were not visible at the surface were marked on SEEG by an epileptologist. Surface signals, MEG and scalp-EEG, were then averaged locked on SEEG spikes. MEG sources were reconstructed based on a moving dipole approach (Brainstorm software). This analysis revealed source within the left occipital pole, located posteriorly to the SEEG leads presenting the maximal number of spikes, in a region not explored by SEEG. In summary, simultaneous recordings provide a new framework for obtaining a view on brain signals that is both local and global, thereby overcoming the inherent SEEG limited spatial sampling.

Response retrieval and motor planning during typing.

Recent work in language production research suggests complex relationships between linguistic and motor processes. Typing is an interesting candidate for investigating further this issue. First, typing presumably relies on the same distributed left-lateralized brain network as handwriting and speech production. Second, typing has its own set of highly specific motor constraints, such as internal keystroke representations that hold information about both letter identity and spatial characteristics of the key to strike. The present study aims to further develop research on typed production, by targeting the dynamics between linguistic and motor neural networks. Specifically, we used a typed picture-naming task to examine the interplay between response retrieval and motor planning. To track processes associated with both linguistic processing and keystroke representation, we manipulated, respectively, the semantic context in which the target appeared and the side of the first keystrokes of the word. We recorded high-density electroencephalography (EEG) continuously from the presentation of a picture, to the typing of its name, and computed both event-related potentials (ERP) and beta-band power analyses. Non-parametric data-driven analysis revealed a clear pattern of response preparation over both hemispheres close to response time, in both the ERP and beta-band power modulations. This was preceded by a left-lateralized power decrease in the beta-band, presumably representing memory retrieval, and an early contrast in ERP, between left and right keystrokes' preparation. We discuss these results in terms of a dynamic access approach for internal keystroke representations, and argue for an integrative rather than separatist view of linguistic and motor processes.

Sequential processing during noun phrase production.

This study examined whether the brain operations involved during the processing of successive words in multi word noun phrase production take place sequentially or simultaneously. German speakers named pictures while ignoring a written distractor superimposed on the picture (picture-word interference paradigm) using the definite determiner and corresponding German noun. The gender congruency and the phonological congruency (i.e., overlap in first phonemes) between target and distractor were manipulated. Naming responses and EEG were recorded. The behavioural performance replicated both the phonology and the gender congruency effects (i.e., shorter naming latencies for gender congruent than incongruent and for phonologically congruent than incongruent trials). The phonological and gender manipulations also influenced the EEG data. Crucially, the two effects occurred in different time windows and over different sets of electrodes. The phonological effect was observed substantially earlier than the gender congruency effect. This finding suggests that the processing of determiners and nouns during determiner noun phrase production occurs at least partly sequentially.

Simultaneous recording of MEG, EEG and intracerebral EEG during visual stimulation: from feasibility to single-trial analysis.

Electroencephalography (EEG), magnetoencephalography (MEG), and intracerebral stereotaxic EEG (SEEG) are the three neurophysiological recording techniques, which are thought to capture the same type of brain activity. Still, the relationships between non-invasive (EEG, MEG) and invasive (SEEG) signals remain to be further investigated. In early attempts at comparing SEEG with either EEG or MEG, the recordings were performed separately for each modality. However such an approach presents substantial limitations in terms of signal analysis. The goal of this technical note is to investigate the feasibility of simultaneously recording these three signal modalities (EEG, MEG and SEEG), and to provide strategies for analyzing this new kind of data. Intracerebral electrodes were implanted in a patient with intractable epilepsy for presurgical evaluation purposes. This patient was presented with a visual stimulation paradigm while the three types of signals were simultaneously recorded. The analysis started with a characterization of the MEG artifact caused by the SEEG equipment. Next, the average evoked activities were computed at the sensor level, and cortical source activations were estimated for both the EEG and MEG recordings; these were shown to be compatible with the spatiotemporal dynamics of the SEEG signals. In the average time-frequency domain, concordant patterns between the MEG/EEG and SEEG recordings were found below the 40 Hz level. Finally, a fine-grained coupling between the amplitudes of the three recording modalities was detected in the time domain, at the level of single evoked responses. Importantly, these correlations have shown a high level of spatial and temporal specificity. These findings provide a case for the ability of trimodal recordings (EEG, MEG, and SEEG) to reach a greater level of specificity in the investigation of brain signals and functions.

Cephalic aura after frontal lobe resection.

A cephalic aura is a common sensory aura typically seen in frontal lobe epilepsy. The generation mechanism of cephalic aura is not fully understood. It is hypothesized that to generate a cephalic aura extensive cortical areas need to be excited. We report a patient who started to have cephalic aura after right frontal lobe resection. Magnetoencephalography (MEG) showed interictal spike and ictal change during cephalic aura, both of which were distributed in the right frontal region, and the latter involved much more widespread areas than the former on MEG sensors. The peculiar seizure onset pattern may indicate that surgical modification of the epileptic network was related to the appearance of cephalic aura. We hypothesize that generation of cephalic aura may be associated with more extensive cortical involvement of epileptic activity than that of interictal activity, in at least a subset of cases.

Magnetoencephalography correlate of EEG POSTS (positive occipital sharp transients of sleep).

PURPOSE: In contrast to EEG, which has guidelines for interpretation and a plethora of textbooks, the full range of activity seen in magnetoencephalography (MEG) has not been fleshed out. Currently, magnetoencephalographers apply criteria for EEG waveforms to MEG signals based on an assumption that MEG activity should have morphology that is similar to EEG. The purpose of this article was to show the characteristic MEG profile of positive occipital sharp transients of sleep. METHODS: Simultaneous MEG-EEG recordings of two cases are shown. RESULTS: In both the cases, the morphologic features of positive occipital sharp transients of sleep in MEG vary and sometimes mimic epileptic spikes. CONCLUSION: This report raises a caution that a normal variant may have an even more epileptic appearance on MEG than on EEG. Using the simultaneously recorded EEG to avoid misinterpretation of spikey-looking positive occipital sharp transients of sleep in MEG is a natural and prudent practice.

Sensitivity of scalp 10-20 EEG and magnetoencephalography.

Although previous studies have investigated the sensitivity of electroencephalography (EEG) and magnetoencephalography (MEG) to detect spikes by comparing simultaneous recordings, there are no published reports that focus on the relationship between spike dipole orientation or sensitivity of scalp EEG/MEG and the "gold standard" of intracranial recording (stereotactic EEG). We evaluated two patients with focal epilepsy; one with lateral temporal focus and the other with insular focus. Two MEG recordings were performed for both patients, each recorded simultaneously with initially scalp EEG, based on international 10-20 electrode placement with additional electrodes for anterior temporal regions, and subsequently stereotactic EEG. Localisation of MEG spike dipoles from both studies was concordant and all MEG spikes were detected by stereotactic EEG. For the patient with lateral temporal epilepsy, spike sensitivity of MEG and scalp EEG (relative to stereotactic EEG) was 55 and 0%, respectively. Of note, in this case, MEG spike dipoles were oriented tangentially to scalp surface in a tight cluster; the angle of the spike dipole to the vertical line was 3.6 degrees. For the patient with insular epilepsy, spike sensitivity of MEG and scalp EEG (relative to stereotactic EEG) was 83 and 44%, respectively; the angle of the spike dipole to the vertical line was 45.3 degrees. For the patient with lateral temporal epilepsy, tangential spikes from the lateral temporal cortex were difficult to detect based on scalp 10-20 EEG and for the patient with insular epilepsy, it was possible to evaluate operculum insular sources using MEG. We believe that these findings may be important for the interpretation of clinical EEG and MEG.

Magnetoencephalography reveals a unique neurophysiological profile of focal-onset epileptic spasms.

Epilepsy is defined as a disorder of the brain characterized by an enduring predisposition to experience epileptic seizures and the neurobiological, cognitive, psychological, and social difficulties relating to the condition. An epileptic spasm (ES) is a type of seizure characterized by clusters of short contractions involving axial muscles and proximal segments. However, the precise mechanism of ESs remains unknown. Despite the potential of magnetoencephalography (MEG) as a tool for investigating the neurophysiological mechanism of ESs, it has been difficult to use this methodology due to magnetic artifacts attributable to patient movement. We report on an 8-year-old girl suffering from intractable epileptic spasms from the age of 7 months. She was diagnosed with possible Aicardi syndrome [corrected] (AGS), characterized by the triad of callosal agenesis, infantile spasms, and chorioretinal lacunae. She is now intellectually delayed and suffers from intractable ES. We used both MEG and electroencephalography to investigate her epilepsy. The recording captured two series of spasm clusters. Spikes were clearly identified with MEG in about four-fifths of all spasms but were identified poorly or not at all in the remainder. MEG findings support previous studies that used intracranial electrodes to analyze patients with ESs and that showed variability in ES-associated spikes in terms of manner of cortical involvement and magnitude. Given the limitations of intracranial electrodes, such as sampling restrictions and invasiveness, MEG may be a helpful tool for non-invasively investigating the unique pathophysiological profile of focal-onset ESs.

Utility of temporally-extended signal space separation algorithm for magnetic noise from vagal nerve stimulators.

OBJECTIVE: To evaluate the utility of a temporally-extended signal space separation algorithm (tSSS) for patients with vagal nerve stimulator (VNS). METHODS: We evaluated median nerve somatosensory evoked responses (SER) of magnetoencephalography (MEG) in 27 VNS patients (48 sides) with/without tSSS processing. We classified SER dipoles as 'acceptable' if: (A) the location of the dipole was in the expected location in the central sulcus, and (B) the goodness of fit value (GOF) was greater than 80%. We evaluated (1) the number of sides which produced acceptable dipoles in each dataset (i.e. with/without tSSS processing), and in cases where the both data produced reliable dipoles, (2) compared their GOFs and the 95% confidence volumes (CV) (mm(3)). Statistical differences in the GOF and CV between with/without tSSS conditions were determined by paired t test. RESULTS: Only 11 (23%) responses had reliable dipoles without tSSS processing, while all 48 (100%) had acceptable dipoles under tSSS processing. Additionally, the latter group had significantly higher GOF (increased by 7% on average) and lower CV (mean decrease of 200 mm(3)) than the former (p<0.01). CONCLUSIONS: Processing with tSSS quantitatively improves dipole fitting of known sources in VNS patients. SIGNIFICANCE: This algorithm permits satisfactory MEG testing in the relatively commonly encountered epilepsy patient with VNS.

Magnetoencephalography in fronto-parietal opercular epilepsy.

OBJECTIVE: To clarify the clinical and neurophysiological profiles of fronto-parietal opercular epilepsy in which epileptic spikes are detected with magnetoencephalography (MEG) but not with scalp electroencephalography (EEG). METHODS: Four patients presented with epileptic spikes localized to the fronto-parietal opercular cortex, which were only appreciated following MEG recordings. RESULTS: In all cases, seizure semiology suggested early activation of the operculum and lower peri-rolandic cortex consistent with the somatotopic organization of this region, i.e. tingling sensation involving the throat and hemi-face or contralateral upper limb, and spasms of the neck and throat. MEG spikes were localized in the fronto-parietal operculum. Three of the four patients underwent invasive electrocorticography and/or stereo-EEG recordings, and spikes were confirmed to arise from the estimated area of MEG dipole localization. Two patients remained seizure-free for over 1 year after resection of the epileptogenic region; the other patient declined resective surgery due to proximity to the language cortex. CONCLUSION: This study demonstrates the usefulness of MEG in localizing spikes arising from within the fronto-parietal opercular regions, and implies that MEG may provide localizing information in patients with symptoms suggestive of opercular epilepsy, even if scalp EEG recordings fail to disclose any epileptogenic activities.

Clinical evidence for the utility of movement compensation algorithm in magnetoencephalography: successful localization during focal seizure.

A movement compensation (MC) algorithm may help to evaluate seizure focus in magnetoencephalography despite patient movement. We report a boy whose ictal MEG focus was localized to the same sublobar region before and after head turning when MC was applied, but which was erroneously localized to a different area without MC. This study provides the first clinical evidence for utility of MC in magnetoencephalography for localizing focal seizures.

Different cortical involvement pattern of generalized and localized spasms: a magnetoencephalography study.

We report successful magnetoencephalography (MEG) recording in a child who had generalized epileptic spasms (ESs) as well as ESs involving the legs only during the recording. MEG source localization results demonstrated that (1) the interictal epileptiform discharges and both types of ESs had the same origin, that is, the right parietal region, and (2) the two types of ESs had different cortical spread patterns, that is, epileptic involvement localized to the right parietal region in spasms of the legs and rapid diffuse involvement in generalized spasms. In this case, MEG provided new insight into the mechanisms underlying the two types of ESs: both types were generated from the same focus, and in generalized ESs, abnormal excitation spread to cortical areas diffusely.

Early brain-body impact of emotional arousal.

Current research in affective neuroscience suggests that the emotional content of visual stimuli activates brain-body responses that could be critical to general health and physical disease. The aim of this study was to develop an integrated neurophysiological approach linking central and peripheral markers of nervous activity during the presentation of natural scenes in order to determine the temporal stages of brain processing related to the bodily impact of emotions. More specifically, whole head magnetoencephalogram (MEG) data and skin conductance response (SCR), a reliable autonomic marker of central activation, were recorded in healthy volunteers during the presentation of emotional (unpleasant and pleasant) and neutral pictures selected from the International Affective Picture System (IAPS). Analyses of event-related magnetic fields (ERFs) revealed greater activity at 180 ms in an occipitotemporal component for emotional pictures than for neutral counterparts. More importantly, these early effects of emotional arousal on cerebral activity were significantly correlated with later increases in SCR magnitude. For the first time, a neuromagnetic cortical component linked to a well-documented marker of bodily arousal expression of emotion, namely, the SCR, was identified and located. This finding sheds light on the time course of the brain-body interaction with emotional arousal and provides new insights into the neural bases of complex and reciprocal mind-body links.

Oscillatory activity in parietal and dorsolateral prefrontal cortex during retention in visual short-term memory: additive effects of spatial attention and memory load.

We used whole-head magnetoencephalography to study the representation of objects in visual short-term memory (VSTM) in the human brain. Subjects remembered the location and color of either two or four colored disks that were encoded from the left or right visual field (equal number of distractors in the other visual hemifield). The data were analyzed using time-frequency methods, which enabled us to discover a strong oscillatory activity in the 8-15 Hz band during the retention interval. The study of the alpha power variation revealed two types of responses, in different brain regions. The first was a decrease in alpha power in parietal cortex, contralateral to the stimuli, with no load effect. The second was an increase of alpha power in parietal and lateral prefrontal cortex, as memory load increased, but without interaction with the hemifield of the encoded stimuli. The absence of interaction between side of encoded stimuli and memory load suggests that these effects reflect distinct underlying mechanisms. A novel method to localize the neural generators of load-related oscillatory activity was devised, using cortically-constrained distributed source-localization methods. Some activations were found in the inferior intraparietal sulcus (IPS) and intraoccipital sulcus (IOS). Importantly, strong oscillatory activity was also found in dorsolateral prefrontal cortex (DLPFC). Alpha oscillatory activity in DLPFC was synchronized with the activity in parietal regions, suggesting that VSTM functions in the human brain may be implemented via a network that includes bilateral DLPFC and bilateral IOS/IPS as key nodes.