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Language is one of the defining abilities of humans. Many studies have characterized the neural correlates of different aspects of language processing. However, the imaging techniques typically used in these studies were limited in either their temporal or spatial resolution. Electrocorticographic (ECoG) recordings from the surface of the brain combine high spatial with high temporal resolution, and thus could be a valuable tool for the study of neural correlates of language function. In this study, we defined the spatiotemporal dynamics of ECoG activity during a word repetition task in nine human subjects. ECoG was recorded while each subject overtly or covertly repeated words that were presented either visually or auditorily. ECoG amplitudes in the high gamma (HG) band confidently tracked neural changes associated with stimulus presentation and with the subject's verbal response. Overt word production was primarily associated with HG changes in the superior and middle parts of temporal lobe, Wernicke's area, the supramarginal gyrus, Broca's area, premotor cortex (PMC), primary motor cortex. Covert word production was primarily associated with HG changes in superior temporal lobe and the supramarginal gyrus. Acoustic processing from both auditory stimuli as well as the subject's own voice resulted in HG power changes in superior temporal lobe and Wernicke's area. In summary, this study represents a comprehensive characterization of overt and covert speech using electrophysiological imaging with high spatial and temporal resolution. It thereby complements the findings of previous neuroimaging studies of language and thus further adds to current understanding of word processing in humans.

The mechanism(s) by which anesthetics reversibly suppress consciousness are incompletely understood. Previous functional imaging studies demonstrated dynamic changes in thalamic and cortical metabolic activity, as well as the maintained presence of metabolically defined functional networks despite the loss of consciousness. However, the invasive electrophysiology associated with these observations has yet to be studied. By recording electrical activity directly from the cortical surface, electrocorticography (ECoG) provides a powerful method to integrate spatial, temporal, and spectral features of cortical electrophysiology not possible with noninvasive approaches. In this study, we report a unique comprehensive recording of invasive human cortical physiology during both induction and emergence from propofol anesthesia. Propofolinduced transitions in and out of consciousness (defined here as responsiveness) were characterized by maintained large-scale functional networks defined by correlated fluctuations of the slow cortical potential (<0.5 Hz) over the somatomotor cortex, present even in the deeply anesthetized state of burst suppression. Similarly, phase-power coupling between θ-and γ-range frequencies persisted throughout the induction and emergence from anesthesia. Superimposed on this preserved functional architecture were alterations in frequency band power, variance, covariance, and phase-power interactions that were distinct to different frequency ranges and occurred in separable phases. These data support that dynamic alterations in cortical and thalamocortical circuit activity occur in the context of a larger stable architecture that is maintained despite anesthetic-induced alterations in consciousness.cortical networks | human cortex | gamma rhythms E very year millions of people undergo general anesthesia, yet the mechanism(s) by which widely used clinical anesthetics reversibly ablate consciousness remains incompletely understood (1). Moreover, the manner in which the brain is able to tolerate global pharmacologic suppression, yet still maintain memories and resume complex cortical interactions that define a person's cognition after removal of this suppression, also remains unknown. Thus far, the majority of studies in humans have used noninvasive methods such as functional imaging and electroencephalography (EEG) to arrive at the current understanding. To date, positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) studies show that there is a complex interplay between and within the thalamus and the cortex. These studies demonstrate that the thalamus is a common site of deactivation during induction by various anesthetic agents (2,3), that there appears to be a disruption of thalamo-cortical and cortico-cortical connectivity (4, 5), and that specific regions of association cortices show enhanced deactivation with certain anesthetics (6, 7). In parallel with these dynamic interactions, there also appear to be physiologic elements that are invariant and do not change wi...

Electrocorticography (ECoG) has emerged as a new signal platform for brain-computer interface (BCI) systems. Classically, the cortical physiology that has been commonly investigated and utilized for device control in humans has been brain signals from sensorimotor cortex. Hence, it was unknown whether other neurophysiological substrates, such as the speech network, could be used to further improve on or complement existing motor-based control paradigms. We demonstrate here for the first time that ECoG signals associated with different overt and imagined phoneme articulation can enable invasively monitored human patients to control a one-dimensional computer cursor rapidly and accurately. This phonetic content was distinguishable within higher gamma frequency oscillations and enabled users to achieve final target accuracies between 68 and 91% within 15 minutes. Additionally, one of the patients achieved robust control using recordings from a microarray consisting of 1 mm spaced microwires. These findings suggest that the cortical network associated with speech could provide an additional cognitive and physiologic substrate for BCI operation and that these signals can be acquired from a cortical array that is small and minimally invasive.

Selective attention allows us to filter out irrelevant information in the environment and focus neural resources on information relevant to our current goals. Functional brain-imaging studies have identified networks of broadly distributed brain regions that are recruited during different attention processes; however, the dynamics by which these networks enable selection are not well understood. Here, we first used functional MRI to localize dorsal and ventral attention networks in human epileptic subjects undergoing seizure monitoring. We subsequently recorded cortical physiology using subdural electrocorticography during a spatialattention task to study network dynamics. Attention networks become selectively phase-modulated at low frequencies (δ, θ) during the same task epochs in which they are recruited in functional MRI. This mechanism may alter the excitability of task-relevant regions or their effective connectivity. Furthermore, different attention processes (holding vs. shifting attention) are associated with synchrony at different frequencies, which may minimize unnecessary cross-talk between separate neuronal processes.O ne of the hallmarks of effective behavior is the ability to flexibly attend to particular stimuli in the environment. Selective attention can be driven endogenously by one's current goals or by salient external stimuli. Human neuroimaging studies have identified two sets of fronto-parietal regions that are recruited during these two types of attention. A set of dorsal fronto-parietal regions (dorsal attention network or DAN) shows sustained activity during endogenous or goal-driven attention (1), and reorienting to unexpected targets transiently activates both the DAN and a second set of regions, the ventral attention network (VAN) (2). Although functional MRI (fMRI) has identified the brain regions that are involved in these attentional operations (3, 4), the slow nature of the hemodynamic response has severely limited the study of network dynamics at behaviorally relevant time scales (5). Here, we report results obtained by cortical surface (electrocorticography or ECoG) recordings in epilepsy patients undergoing clinical monitoring to identify seizure foci. Electrode locations for each subject were colocalized with functional brain networks, including the DAN and VAN identified in the same subjects using fMRI. This experimental paradigm allowed us to objectively link fast electrophysiological dynamics, during performance of an attention task, to well-studied functional brain networks.ECoG measures nonspiking, local field potential oscillations across a range of frequencies, which are thought to reflect fluctuations in local neuronal excitability (6, 7). Phase modulations of activity within a region and between regions may therefore affect, respectively, their ability to respond to inputs and to transfer information between one another (8, 9). In support of this theory, previous studies in both animals and humans have shown that either local or long-distance synchrony change in a tas...

High-gamma band (>60Hz) power changes in cortical electrophysiology are a reliable indicator of focal, event-related cortical activity. In spite of discoveries of oscillatory subthreshold and synchronous suprathreshold activity at the cellular level, there is an increasingly popular view that high-gamma band amplitude changes recorded from cellular ensembles are the result of asynchronous firing activity that yields wideband and uniform power increases. Others have demonstrated independence of power changes in the low- and high-gamma bands, but to date, no studies have shown evidence of any such independence above 60Hz. Based on non-uniformities in time-frequency analyses of electrocorticographic (ECoG) signals, we hypothesized that induced high-gamma band (60-500Hz) power changes are more heterogeneous than currently understood. Using single-word repetition tasks in six human subjects, we showed that functional responsiveness of different ECoG high-gamma sub-bands can discriminate cognitive task (e.g., hearing, reading, speaking) and cortical locations. Power changes in these sub-bands of the high-gamma range are consistently present within single trials and have statistically different time courses within the trial structure. Moreover, when consolidated across all subjects within three task-relevant anatomic regions (sensorimotor, Broca's area, and superior temporal gyrus), these behavior- and location- dependent power changes evidenced nonuniform trends across the population. Taken together, the independence and nonuniformity of power changes across a broad range of frequencies suggest that a new approach to evaluating high-gamma band cortical activity is necessary. These findings show that in addition to time and location, frequency is another fundamental dimension of high-gamma dynamics.

Several scientists have proposed different models for cortical processing of speech. Classically, the regions participating in language were thought to be modular with a linear sequence of activations. More recently, modern theoretical models have posited a more hierarchical and distributed interaction of anatomic areas for the various stages of speech processing. Traditional imaging techniques can only define the location or time of cortical activation, which impedes the further evaluation and refinement of these models. In this study, we take advantage of recordings from the surface of the brain [electrocorticography (ECoG)], which can accurately detect the location and timing of cortical activations, to study the time course of ECoG high gamma (HG) modulations during an overt and covert word repetition task for different cortical areas. For overt word production, our results show substantial perisylvian cortical activations early in the perceptual phase of the task that were maintained through word articulation. However, this broad activation is attenuated during the expressive phase of covert word repetition. Across the different repetition tasks, the utilization of the different cortical sites within the perisylvian region varied in the degree of activation dependent on which stimulus was provided (auditory or visual cue) and whether the word was to be spoken or imagined. Taken together, the data support current models of speech that have been previously described with functional imaging. Moreover, this study demonstrates that the broad perisylvian speech network activates early and maintains suprathreshold activation throughout the word repetition task that appears to be modulated by the demands of different conditions.

Objective Electrocorticography (ECoG) electrodes implanted on the surface of the brain have recently emerged as a potential signal platform for brain-computer interface (BCI) systems. While clinical ECoG electrodes are currently implanted beneath the dura, epidural electrodes could reduce the invasiveness and the potential impact of a surgical site infection. Subdural electrodes, on the other hand, while slightly more invasive, may have better signals for BCI application. Because of this balance between risk and benefit between the two electrode positions, the effect of the dura on signal quality must be determined in order to define the optimal implementation for an ECoG BCI system. Approach This study utilized simultaneously acquired baseline recordings from epidural and subdural ECoG electrodes while patients rested. Both macro-scale (2 mm diameter electrodes with 1 cm inter-electrode distance, 1 patient) and micro-scale (75 μm diameter electrodes with 1 mm inter-electrode distance, 4 patients) ECoG electrodes were tested. Signal characteristics were evaluated to determine differences in the spectral amplitude and noise floor. Furthermore, the experimental results were compared to theoretical effects produced by placing epidural and subdural ECoG contacts of different sizes within a finite element model. Main Results The analysis demonstrated that for micro-scale electrodes, subdural contacts have significantly higher spectral amplitudes and reach the noise floor at a higher frequency than epidural contacts. For macro-scale electrodes, while there are statistical differences, these differences are small in amplitude and likely do not represent differences relevant to the ability of the signals to be used in a BCI system. Conclusions Our findings demonstrate an important trade-off that should be considered in developing a chronic BCI system. While implanting electrodes under the dura is more invasive, it is associated with increased signal quality when recording from micro-scale electrodes with very small sizes and spacing. If recording from larger electrodes, such as traditionally used clinically, the signal quality of epidural recordings is similar to that of subdural recordings.