Sarang S. Dalal

High Gamma Power Is Phase-Locked to Theta Oscillations in Human Neocortex

We observed robust coupling between the high- and low-frequency bands of ongoing electrical activity in the human brain. In particular, the phase of the low-frequency theta (4 to 8 hertz) rhythm modulates power in the high gamma (80 to 150 hertz) band of the electrocorticogram, with stronger modulation occurring at higher theta amplitudes. Furthermore, different behavioral tasks evoke distinct patterns of theta/high gamma coupling across the cortex. The results indicate that transient coupling between low- and high-frequency brain rhythms coordinates activity in distributed cortical areas, providing a mechanism for effective communication during cognitive processing in humans.

Five-dimensional neuroimaging : localization of the time–frequency dynamics of cortical activity

The spatiotemporal dynamics of cortical oscillations across human brain regions remain poorly understood because of a lack of adequately validated methods for reconstructing such activity from noninvasive electrophysiological data. In this paper, we present a novel adaptive spatial filtering algorithm optimized for robust source time-frequency reconstruction from magnetoencephalography (MEG) and electroencephalography (EEG) data. The efficacy of the method is demonstrated with simulated sources and is also applied to real MEG data from a self-paced finger movement task. The algorithm reliably reveals modulations both in the beta band (12-30 Hz) and high gamma band (65-90 Hz) in sensorimotor cortex. The performance is validated by both across-subjects statistical comparisons and by intracranial electrocorticography (ECoG) data from two epilepsy patients. Interestingly, we also reliably observed high frequency activity (30-300 Hz) in the cerebellum, although with variable locations and frequencies across subjects. The proposed algorithm is highly parallelizable and runs efficiently on modern high-performance computing clusters. This method enables the ultimate promise of MEG and EEG for five-dimensional imaging of space, time, and frequency activity in the brain and renders it applicable for widespread studies of human cortical dynamics during cognition.

Task-related gamma-band dynamics from an intracerebral perspective: review and implications for surface EEG and MEG.

Although non‐invasive techniques provide functional activation maps at ever‐growing spatio‐temporal precision, invasive recordings offer a unique opportunity for direct investigations of the fine‐scale properties of neural mechanisms in focal neuronal populations. In this review we provide an overview of the field of intracranial Electroencephalography (iEEG) and discuss its strengths and limitations and its relationship to non‐invasive brain mapping techniques. We discuss the characteristics of invasive data acquired from implanted epilepsy patients using stereotactic‐electroencephalography (SEEG) and electrocorticography (ECoG) and the use of spectral analysis to reveal task‐related modulations in multiple frequency components. Increasing evidence suggests that gamma‐band activity (>40 Hz) might be a particularly efficient index for functional mapping. Moreover, the detection of high gamma activity may play a crucial role in bridging the gap between electrophysiology and functional imaging studies as well as in linking animal and human data. The present review also describes recent advances in real‐time invasive detection of oscillatory modulations (including gamma activity) in humans. Furthermore, the implications of intracerebral findings on future non‐invasive studies are discussed. Hum Brain Mapp, 2009.

Spatiotemporal dynamics of word processing in the human brain

We examined the spatiotemporal dynamics of word processing by recording the electrocorticogram (ECoG) from the lateral frontotemporal cortex of neurosurgical patients chronically implanted with subdural electrode grids. Subjects engaged in a target detection task where proper names served as infrequent targets embedded in a stream of task-irrelevant verbs and nonwords. Verbs described actions related to the hand (e.g, throw) or mouth (e.g., blow), while unintelligible nonwords were sounds which matched the verbs in duration, intensity, temporal modulation, and power spectrum. Complex oscillatory dynamics were observed in the delta, theta, alpha, beta, low, and high gamma (HG) bands in response to presentation of all stimulus types. HG activity (80–200 Hz) in the ECoG tracked the spatiotemporal dynamics of word processing and identified a network of cortical structures involved in early word processing. HG was used to determine the relative onset, peak, and offset times of local cortical activation during word processing. Listening to verbs compared to nonwords sequentially activates first the posterior superior temporal gyrus (post-STG), then the middle superior temporal gyrus (mid-STG), followed by the superior temporal sulcus (STS). We also observed strong phase-locking between pairs of electrodes in the theta band, with weaker phase-locking occurring in the delta, alpha, and beta frequency ranges. These results provide details on the first few hundred milliseconds of the spatiotemporal evolution of cortical activity during word processing and provide evidence consistent with the hypothesis that an oscillatory hierarchy coordinates the flow of information between distinct cortical regions during goal-directed behavior.

Mapping Functional Connectivity in Patients with Brain Lesions

The spatial distribution of functional connectivity between brain areas and the disturbance introduced by focal brain lesions are poorly understood. Based on the rationale that damaged brain tissue is disconnected from the physiological interactions among healthy areas, this study aimed to map the functionality of brain areas according to their connectivity with other areas.

Modified beamformers for coherent source region suppression

Many tomographic source localization algorithms used in biomagnetic imaging assume, explicitly or sometimes implicitly, that the source activity at different brain locations are either independent or that the correlation structure between sources is known. Among these algorithms is a class of adaptive spatial filters known as beamformers, which have superior spatiotemporal resolution abilities. The performance of beamformers is robust to weakly coherent sources. However, these algorithms are extremely sensitive to the presence of strongly coherent sources. A frequent mode of failure in beamformers occurs with reconstruction of auditory evoked fields (AEFs), in which bilateral auditory cortices are highly coherent in their activation. Here, we present a novel beamformer that suppresses activation from regions with interfering coherent sources. First, a volume containing the interfering sources is defined. The lead field matrix for this volume is computed and reduced into a few significant columns using singular value decomposition (SVD). A vector beamformer is then constructed by rejecting the contribution of sources in the suppression region while allowing for source reconstruction at other specified regions. Performance of this algorithm was first validated with simulated data. Subsequent tests of this modified beamformer were performed on bilateral AEF data. An unmodified vector beamformer using whole head coverage misplaces the source medially. After defining a suppression region containing the temporal cortex on one side, the described method consistently results in clear focal activations at expected regions of the contralateral superior temporal plane.

Comparison of Time–Frequency Responses and the Event-Related Potential to Auditory Speech Stimuli in Human Cortex

We recorded the electrocorticogram directly from the exposed cortical surface of awake neurosurgical patients during the presentation of auditory syllable stimuli. All patients were unanesthetized as part of a language-mapping procedure for subsequent left-hemisphere tumor resection. Time-frequency analyses showed significant high-gamma (gammahigh: 70-160 Hz) responses from the left superior temporal gyrus, but no reliable response from the left inferior frontal gyrus. Alpha suppression (alpha: 7-14 Hz) and event-related potential responses exhibited a more widespread topography. Across electrodes, the alpha suppression from 200 to 450 ms correlated with the preceding (50-200 ms) gammahigh increase. The results are discussed in terms of the different physiological origins of these electrocortical signals.

Exploring the electrophysiological correlates of the default-mode network with intracerebral EEG

While functional imaging studies allow for a precise spatial characterization of resting state networks, their neural correlates and thereby their fine-scale temporal dynamics remain elusive. A full understanding of the mechanisms at play requires input from electrophysiological studies. Here, we discuss human and non-human primate electrophysiological data that explore the neural correlates of the default-mode network. Beyond the promising findings obtained with non-invasive approaches, emerging evidence suggests that invasive recordings in humans will be crucial in order to elucidate the neural correlates of the brains default-mode function. In particular, we contend that stereotactic-electroencephalography, which consists of implanting multiple depth electrodes for pre-surgical evaluation in drug-resistant epilepsy, is particularly suited for this endeavor. We support this view by providing rare data from depth recordings in human posterior cingulate cortex and medial prefrontal cortex that show transient neural deactivation during task-engagement.

Simultaneous MEG and intracranial EEG recordings during attentive reading

The relationship between neural oscillations recorded at various spatial scales remains poorly understood partly due to an overall dearth of studies utilizing simultaneous measurements. In an effort to study quantitative markers of attention during reading, we performed simultaneous magnetoencephalography (MEG) and intracranial electroencephalography (iEEG) recordings in four epileptic patients. Patients were asked to attend to a specific color when presented with an intermixed series of red words and green words, with words of a given color forming a cohesive story. We analyzed alpha, beta, and gamma band oscillatory responses to the word presentation and compared the strength and spatial organization of those responses in both electrophysiological recordings. Time-frequency analysis of iEEG revealed a network of clear attention-modulated high gamma band (50-150 Hz) power increases and alpha/beta (9-25 Hz) suppressions in response to the words. In addition to analyses at the sensor level, MEG time-frequency analysis was performed at the source level using a sliding window beamformer technique. Strong alpha/beta suppressions were observed in MEG reconstructions, in tandem with iEEG effects. While the MEG counterpart of high gamma band enhancement was difficult to interpret at the sensor level in two patients, MEG time-frequency source reconstruction revealed additional activation patterns in accordance with iEEG results. Importantly, iEEG allowed us to confirm that several sources of gamma band modulation observed with MEG were indeed of cortical origin rather than EMG muscular or ocular artifact.

Category-Specific Visual Responses: An Intracranial Study Comparing Gamma, Beta, Alpha, and ERP Response Selectivity.

The specificity of neural responses to visual objects is a major topic in visual neuroscience. In humans, functional magnetic resonance imaging (fMRI) studies have identified several regions of the occipital and temporal lobe that appear specific to faces, letter strings, scenes, or tools. Direct electrophysiological recordings in the visual cortical areas of epileptic patients have largely confirmed this modular organization, using either single-neuron peri-stimulus time-histogram or intracerebral event-related potentials (iERP). In parallel, a new research stream has emerged using high-frequency gamma-band activity (50–150 Hz) (GBR) and low-frequency alpha/beta activity (8–24 Hz) (ABR) to map functional networks in humans. An obvious question is now whether the functional organization of the visual cortex revealed by fMRI, ERP, GBR, and ABR coincide. We used direct intracerebral recordings in 18 epileptic patients to directly compare GBR, ABR, and ERP elicited by the presentation of seven major visual object categories (faces, scenes, houses, consonants, pseudowords, tools, and animals), in relation to previous fMRI studies. Remarkably both GBR and iERP showed strong category-specificity that was in many cases sufficient to infer stimulus object category from the neural response at single-trial level. However, we also found a strong discrepancy between the selectivity of GBR, ABR, and ERP with less than 10% of spatial overlap between sites eliciting the same category-specificity. Overall, we found that selective neural responses to visual objects were broadly distributed in the brain with a prominent spatial cluster located in the posterior temporal cortex. Moreover, the different neural markers (GBR, ABR, and iERP) that elicit selectivity toward specific visual object categories present little spatial overlap suggesting that the information content of each marker can uniquely characterize high-level visual information in the brain.