Language processing requires the orchestrated action of different neuronal populations, and some studies suggest that the role of the basal temporal (BT) cortex in language processing is bilaterally distributed. Our aim was to demonstrate connectivity between perisylvian cortex and both BT areas. We recorded corticocortical evoked potentials (CCEPs) in 8 patients with subdural electrodes implanted for surgical evaluation of intractable epilepsy. Four patients had subdural grids over dominant perisylvian and BT areas, and 4 had electrode strips over both BT areas and left posterior superior temporal gyrus (LPSTG). After electrocortical mapping, patients with grids had 1-Hz stimulation of language areas. Patients with strips did not undergo mapping but had 1-Hz stimulation of the LPSTG. Posterior language area stimulation elicited CCEPs in ipsilateral BT cortex in 3/4 patients with left hemispheric grids. CCEPs were recorded in bilateral BT cortices in 3/4 patients with strips upon stimulation of the LPSTG, and in the LPSTG in the fourth patient upon stimulation of either BT area. This is the first in vivo demonstration of connectivity between LPSTG and both BT cortices. The role of BT cortex in language processing may be bilaterally distributed and related to linking visual information with phonological representations stored in the LPSTG.
basal temporal language area; connectivity; cortical mapping; epilepsy; evoked potentials
There is considerable lay discussion that children with Attention-Deficit/Hyperactivity Disorder (ADHD) have increased difficult with multitasking, but there are few experimental data. In the current study, we examine the simultaneous processing of two stimulus-response tasks using the psychological refractory period (PRP) effect. We hypothesized that children with ADHD would show a greater PRP effect, suggesting a prolonged “bottleneck” in stimulus-response processing. A total of 19 school-aged children with ADHD showed a prolonged PRP effect compared with 25 control children, suggesting a higher cognitive cost in ADHD for multi-tasking.
Multi-tasking; Executive Function; ADHD; Processing Speed; Psychological Refractory Period
Intracranial EEG studies in humans have shown that functional brain activation in a variety of functional-anatomic domains of human cortex is associated with an increase in power at a broad range of high gamma (> 60 Hz) frequencies. Although these electrophysiological responses are highly specific for the location and timing of cortical processing and in animal recordings are highly correlated with increased population firing rates, there has been little direct empirical evidence for causal interactions between different recording sites at high gamma frequencies. Such causal interactions are hypothesized to occur during cognitive tasks that activate multiple brain regions. To determine whether such causal interactions occur at high gamma frequencies and to investigate their functional significance, we used event-related causality (ERC) analysis to estimate the dynamics, directionality, and magnitude of event-related causal interactions using subdural electrocorticography (ECoG) recorded during two word production tasks: picture naming and auditory word repetition. A clinical subject who had normal hearing but was skilled in American Signed Language (ASL) provided a unique opportunity to test our hypothesis with reference to a predictable pattern of causal interactions, i.e. that language cortex interacts with different areas of sensorimotor cortex during spoken vs. signed responses. Our ERC analyses confirmed this prediction. During word production with spoken responses, perisylvian language sites had prominent causal interactions with mouth/tongue areas of motor cortex, and when responses were gestured in sign language, the most prominent interactions involved hand and arm areas of motor cortex. Furthermore, we found that the sites from which the most numerous and prominent causal interactions originated, i.e. sites with a pattern of ERC “divergence”, were also sites where high gamma power increases were most prominent and where electrocortical stimulation mapping interfered with word production. These findings suggest that the number, strength and directionality of event-related causal interactions may help identify network nodes that are not only activated by a task but are critical to its performance.
effective connectivity; Granger causality; large-scale brain networks; high gamma oscillations; language mapping
Recent studies in primate neurophysiology have focused on decoding multi-joint kinematics from single unit and local field potential recordings. However, the extent to which these results can be generalized to human subjects is not known. We have recorded simultaneous electrocorticographic (ECoG) and hand kinematics in a human subject performing reach-grasp-hold of objects varying in shape and size. All Spectral features in various gamma bands (30–50 Hz, 70–100 Hz and 100–150 Hz frequency bands) were able to predict the time course of grasp aperture with high correlation (max r = 0.80) using as few as one ECoG feature from a single electrode (max r for single feature = 0.75) in single trials without prior knowledge of task timing. These results suggest that the population activity captured with ECoG contains information about coordinated finger movements that potentially can be exploited to control advanced upper limb neuroprosthetics.
Recent work has shown that large amplitude negative periods in the local field potential (nLFPs) are able to spread in saltatory manner across large distances in the cortex without distortion in their temporal structure forming ‘coherence potentials’. Here we analysed subdural electrocorticographic (ECoG) signals recorded at 59 sites in the sensorimotor cortex in the left hemisphere of a human subject performing a simple visuomotor task (fist clenching and foot dorsiflexion) to understand how coherence potentials arising in the recordings relate to sensorimotor behavior. In all behaviors we found a particular coherence potential (i.e. a cascade of a particular nLFP wave pattern) arose consistently across all trials with temporal specificity. During contrateral fist clenching, but not the foot dorsiflexion or ipsilateral fist clenching, the coherence potential most frequently originated in the hand representation area in the somatosensory cortex during the anticipation and planning periods of the trial, moving to other regions during the actual motor behavior. While these ‘expert’ sites participated more consistently, other sites participated only a small fraction of the time. Furthermore, the timing of the coherence potential at the hand representation area after onset of the cue predicted the timing of motor behavior. We present the hypothesis that coherence potentials encode information relevant for behavior and are generated by the ‘expert’ sites that subsequently broadcast to other sites as a means of ‘sharing knowledge’.
Direct brain recordings from neurosurgical patients listening to speech reveal that the acoustic speech signals can be reconstructed from neural activity in auditory cortex.
How the human auditory system extracts perceptually relevant acoustic features of speech is unknown. To address this question, we used intracranial recordings from nonprimary auditory cortex in the human superior temporal gyrus to determine what acoustic information in speech sounds can be reconstructed from population neural activity. We found that slow and intermediate temporal fluctuations, such as those corresponding to syllable rate, were accurately reconstructed using a linear model based on the auditory spectrogram. However, reconstruction of fast temporal fluctuations, such as syllable onsets and offsets, required a nonlinear sound representation based on temporal modulation energy. Reconstruction accuracy was highest within the range of spectro-temporal fluctuations that have been found to be critical for speech intelligibility. The decoded speech representations allowed readout and identification of individual words directly from brain activity during single trial sound presentations. These findings reveal neural encoding mechanisms of speech acoustic parameters in higher order human auditory cortex.
Spoken language is a uniquely human trait. The human brain has evolved computational mechanisms that decode highly variable acoustic inputs into meaningful elements of language such as phonemes and words. Unraveling these decoding mechanisms in humans has proven difficult, because invasive recording of cortical activity is usually not possible. In this study, we take advantage of rare neurosurgical procedures for the treatment of epilepsy, in which neural activity is measured directly from the cortical surface and therefore provides a unique opportunity for characterizing how the human brain performs speech recognition. Using these recordings, we asked what aspects of speech sounds could be reconstructed, or decoded, from higher order brain areas in the human auditory system. We found that continuous auditory representations, for example the speech spectrogram, could be accurately reconstructed from measured neural signals. Reconstruction quality was highest for sound features most critical to speech intelligibility and allowed decoding of individual spoken words. The results provide insights into higher order neural speech processing and suggest it may be possible to readout intended speech directly from brain activity.
Subdural electrodes are frequently used to aid in the neurophysiological assessment of patients with intractable seizures. We review the indications for these, their uses for localizing epileptogenic regions and for localizing cortical regions supporting movement, sensation, and language.
Subdural electrodes; implanted electrodes; intracranial EEG recordings; functional localization
Multilingual patients pose a unique challenge when planning epilepsy surgery near language cortex because the cortical representations of each language may be distinct. These distinctions may not be evident with routine electrocortical stimulation mapping (ESM). Electrocorticography (ECoG) has recently been used to detect task-related spectral perturbations associated with functional brain activation. We hypothesized that using broadband high gamma augmentation (HGA, 60–150 Hz) as an index of cortical activation, ECoG would complement ESM in discriminating the cortical representations of first (L1) and second (L2) languages. We studied four adult patients for whom English was a second language, in whom subdural electrodes (a total of 358) were implanted to guide epilepsy surgery. Patients underwent ECoG recordings and ESM while performing the same visual object naming task in L1 and L2. In three of four patients, ECoG found sites activated during naming in one language but not the other. These language-specific sites were not identified using ESM. In addition, ECoG HGA was observed at more sites during L2 versus L1 naming in two patients, suggesting that L2 processing required additional cortical resources compared to L1 processing in these individuals. Post-operative language deficits were identified in three patients (one in L2 only). These deficits were predicted by ECoG spectral mapping but not by ESM. These results suggest that pre-surgical mapping should include evaluation of all utilized languages to avoid post-operative functional deficits. Finally, this study suggests that ECoG spectral mapping may potentially complement the results of ESM of language.
multilingual; naming; electrocorticography; ECoG; high gamma; functional mapping; electrocortical stimulation mapping; epilepsy surgery
Many infants born with a facial port-wine (PW) birthmark will not develop brain involvement of Sturge-Weber syndrome (SWS). Previous studies have shown asymmetry in quantitative EEG (qEEG) correlates with degree of clinical impairment in children and adults with known SWS. We hope to determine if quantitative qEEG can be used as a method to predict which infants are most likely to develop SWS brain involvement on MRI. The current study looks at the ability of qEEG to differentiate between infants with radiographically-demonstrated SWS and those without.
We first performed an observational study of qEEG results on 8 infants with facial PW birthmark (4 had SWS brain involvement). We recorded standard clinical EEGs and then derived a measure of asymmetry. We subsequently validated this threshold through a study of an additional 9 infants with PW birthmark (5 with SWS brain involvement).
Quantitative EEG correctly identified infants with SWS brain involvement in all cases in the validation cohort. This technique was at least as good as a pediatric electroencephalographer with extensive experience reading SWS EEGs.
This study demonstrates the ability for qEEG to discriminate between those infants with SWS brain involvement and those with neurologically asymptomatic PW birthmark.
This study represents an important step toward the development of a qEEG technique able to predict which infants with PW birthmark will develop SWS brain involvement.
Sturge-Weber syndrome; electroencephalography
Multichannel intracranial recordings are used increasingly to study the functional organization of human cortex. Intracranial recordings of event-related activity, or electrocorticography (ECoG), are based on high density electrode arrays implanted directly over cortex, combining good temporal and spatial resolution. Developing appropriate statistical methods for analyzing event-related responses in these high dimensional ECoG datasets remains a major challenge for clinical and systems neuroscience. We present a novel methodological framework that combines complementary, existing methods adapted for statistical analysis of auditory event-related responses in multichannel ECoG recordings. This analytic framework integrates single-channel (time-domain, time–frequency) and multichannel analyses of event-related ECoG activity to determine statistically significant evoked responses, induced spectral responses, and effective (causal) connectivity. Implementation of this quantitative approach is illustrated using multichannel ECoG data from recent studies of auditory processing in patients with epilepsy. Methods described include a time–frequency matching pursuit algorithm adapted for modeling brief, transient cortical spectral responses to sound, and a recently developed method for estimating effective connectivity using multivariate autoregressive modeling to measure brief event-related changes in multichannel functional interactions. A semi-automated spatial normalization method for comparing intracranial electrode locations across patients is also described. The individual methods presented are published and readily accessible. We discuss the benefits of integrating multiple complementary methods in a unified and comprehensive quantitative approach. Methodological considerations in the analysis of multichannel ECoG data, including corrections for multiple comparisons are discussed, as well as remaining challenges in the development of new statistical approaches.
electrocorticography; auditory; matching pursuit; multivariate autoregressive modeling; epilepsy; statistical testing
The phase of ongoing theta (4–8 Hz) and alpha (8–12 Hz) electrophysiological oscillations is coupled to high gamma (80–150 Hz) amplitude, which suggests that low-frequency oscillations modulate local cortical activity. While this phase–amplitude coupling (PAC) has been demonstrated in a variety of tasks and cortical regions, it has not been shown whether task demands differentially affect the regional distribution of the preferred low-frequency coupling to high gamma. To address this issue we investigated multiple-rhythm theta/alpha to high gamma PAC in two subjects with implanted subdural electrocorticographic grids. We show that high gamma amplitude couples to the theta and alpha troughs and demonstrate that, during visual tasks, alpha/high gamma coupling preferentially increases in visual cortical regions. These results suggest that low-frequency phase to high-frequency amplitude coupling is modulated by behavioral task and may reflect a mechanism for selection between communicating neuronal networks.
electrocorticogram; oscillations; cross-frequency coupling; phase–amplitude coupling; gamma; alpha; theta
Recent studies using electrocorticographic (ECoG) recordings in humans have shown that functional activation of cortex is associated with an increase in power in the high-gamma frequency range (∼60–200 Hz). Here we investigate the neural correlates of this high-gamma activity in local field potential (LFP). Single units and LFP were recorded with microelectrodes from the hand region of macaque SII cortex while vibrotactile stimuli of varying intensities were presented to the hand. We found that high-gamma power in the LFP was strongly correlated with the average firing rate recorded by the microelectrodes, both temporally and on a trial-by-trial basis. In comparison, the correlation between firing rate and low-gamma power (40–80 Hz) was much smaller. In order to explore the potential effects of neuronal firing on ECoG, we developed a model to estimate ECoG power generated by different firing patterns of the underlying cortical population and studied how ECoG power varies with changes in firing rate versus the degree of synchronous firing between neurons in the population. Both an increase in firing rate and neuronal synchrony increased high-gamma power in the simulated ECoG data. However, ECoG high-gamma activity was much more sensitive to increases in neuronal synchrony than firing rate.
Secondary somatosensory cortex; gamma; high-gamma; local field potential; ECoG; synchrony
Neuronal oscillations in the gamma frequency range have been reported in many cortical areas, but the role they play in cortical processing remains unclear. We tested a recently proposed hypothesis that the intensity of sensory input is coded in the timing of action potentials relative to the phase of gamma oscillations, thus converting amplitude information to a temporal code. We recorded spikes and local field potential (LFP) from secondary somatosensory (SII) cortex in awake monkeys while presenting a vibratory stimulus at different amplitudes. We developed a novel technique based on matching pursuit to study the interaction between the highly transient gamma oscillations and spikes with high time-frequency resolution. We found that spikes were weakly coupled to LFP oscillations in the gamma frequency range (40−80 Hz), and strongly coupled to oscillations in higher gamma frequencies. However, the phase relationship of neither low-gamma nor high-gamma oscillations changed with stimulus intensity, even with a ten-fold increase. We conclude that, in SII, gamma oscillations are synchronized with spikes, but their phase does not vary with stimulus intensity. Furthermore, high-gamma oscillations (>60 Hz) appear to be closely linked to the occurrence of action potentials, suggesting that LFP high-gamma power could be a sensitive index of the population firing rate near the microelectrode.
Secondary somatosensory cortex; gamma; high-gamma; phase coding; local field potential; matching pursuit
Patterns of responses in the cerebral cortex can vary, and are influenced by pre-existing cortical function, but it is not known how rapidly these variations can occur in humans. We investigated how rapidly response patterns to electrical stimulation can vary in intact human brain. We also investigated whether the type of functional change occurring at a given location with stimulation would help predict the distribution of responses elsewhere over the cortex to stimulation at that given location. We did this by studying cortical afterdischarges following electrical stimulation of the cortex in awake humans undergoing evaluations for brain surgery. Response occurrence and location could change within seconds, both nearby to and distant from stimulation sites. Responses might occur at a given location during one trial but not the next. They could occur at electrodes adjacent or not adjacent to those directly stimulated or to other electrodes showing afterdischarges. The likelihood of an afterdischarge at an individual site after stimulation was predicted by spontaneous electroencephalographic activity at that specific site just prior to stimulation, but not by overall cortical activity. When stimulation at a site interrupted motor, sensory or language function, afterdischarges were more likely to occur at other sites where stimulation interrupted similar functions. These results show that widespread dynamic changes in cortical responses can occur in intact cortex within short periods of time, and that the distribution of these responses depends on local brain states and functional brain architecture at the time of stimulation. Similar rapid variations may occur during normal intracortical communication and may underlie changes in the cortical organization of function. Possibly these variations, and the occurrence and distribution of responses to cortical stimulation, could be predicted. If so, interventions such as stimulation might be used to alter spread of epileptogenic activity, accelerate learning or enhance cortical reorganization after brain injury.
electrical stimulation; brain activation; brain circuits; cerebral electrophysiology; epileptiform EEG discharges
The use of neural signals for prosthesis control is an emerging frontier of research to restore lost function to amputees and the paralyzed. Electrocorticography (ECoG) brain-machine interfaces (BMI) are an alternative to EEG and neural spiking and local field potential BMI approaches. Conventional ECoG BMIs rely on spectral analysis at specific electrode sites to extract signals for controlling prostheses. We compare traditional features with information about the connectivity of an ECoG electrode network. We use time-varying dynamic Bayesian networks (TV-DBN) to determine connectivity between ECoG channels in humans during a motor task. We show that, on average, TV-DBN connectivity decreases from baseline preceding movement and then becomes negative, indicating an alteration in the phase relationship between electrode pairs. In some subjects, this change occurs preceding and during movement, before changes in low or high frequency power. We tested TV-DBN output in a hand kinematic decoder and obtained an average correlation coefficient (r2) between actual and predicted joint angle of 0.40, and as high as 0.66 in one subject. This result compares favorably with spectral feature decoders, for which the average correlation coefficient was 0.13. This work introduces a new feature set based on connectivity and demonstrates its potential to improve ECoG BMI accuracy.
Brain computer interfaces; connectivity analysis; motor control; time-varying dynamic Bayesian networks
To study the role of gamma oscillations (>30 Hz) in selective attention using subdural electrocorticography (ECoG) in humans.
We recorded ECoG in human subjects implanted with subdural electrodes for epilepsy surgery. Sequences of auditory tones and tactile vibrations of 800 ms duration were presented asynchronously, and subjects were asked to selectively attend to one of the two stimulus modalities in order to detect an amplitude increase at 400 ms in some of the stimuli.
Event-related ECoG gamma activity was greater over auditory cortex when subjects attended auditory stimuli and was greater over somatosensory cortex when subjects attended vibrotactile stimuli. Furthermore, gamma activity was also observed over prefrontal cortex when stimuli appeared in either modality, but only when they were attended. Attentional modulation of gamma power began ∼400 ms after stimulus onset, consistent with the temporal demands on attention. The increase in gamma activity was greatest at frequencies between 80 and 150 Hz, in the so-called high gamma frequency range.
There appears to be a strong link between activity in the high-gamma range (80-150 Hz) and selective attention.
Selective attention is correlated with increased activity in a frequency range that is significantly higher than what has been reported previously using EEG recordings.
Attention; ECoG; Gamma oscillations; High-Gamma Activity; Sensory cortex; Intracranial EEG