Spatiotemporal dynamics of bimanual integration in human somatosensory cortex and their relevance to bimanual object manipulation.

Little is known about the spatiotemporal dynamics of cortical responses that integrate slightly asynchronous somatosensory inputs from both hands. This study aimed to clarify the timing and magnitude of interhemispheric interactions during early integration of bimanual somatosensory information in different somatosensory regions and their relevance for bimanual object manipulation and exploration. Using multi-fiber probabilistic diffusion tractography and MEG source analysis of conditioning-test (C-T) median nerve somatosensory evoked fields in healthy human subjects, we sought to extract measures of structural and effective callosal connectivity between different somatosensory cortical regions and correlated them with bimanual tactile task performance. Neuromagnetic responses were found in major somatosensory regions, i.e., primary somatosensory cortex SI, secondary somatosensory cortex SII, posterior parietal cortex, and premotor cortex. Contralateral to the test stimulus, SII activity was maximally suppressed by 51% at C-T intervals of 40 and 60 ms. This interhemispheric inhibition of the contralateral SII source activity correlated directly and topographically specifically with the fractional anisotropy of callosal fibers interconnecting SII. Thus, the putative pathway that mediated inhibitory interhemispheric interactions in SII was a transcallosal route from ipsilateral to contralateral SII. Moreover, interhemispheric inhibition of SII source activity correlated directly with bimanual tactile task performance. These findings were exclusive to SII. Our data suggest that early interhemispheric somatosensory integration primarily occurs in SII, is mediated by callosal fibers that interconnect homologous SII areas, and has behavioral importance for bimanual object manipulation and exploration.

Spontaneous and visually driven high-frequency oscillations in the occipital cortex: intracranial recording in epileptic patients.

High-frequency oscillations (HFOs) at ≥80 Hz of nonepileptic nature spontaneously emerge from human cerebral cortex. In 10 patients with extraoccipital lobe epilepsy, we compared the spectral-spatial characteristics of HFOs spontaneously arising from the nonepileptic occipital cortex with those of HFOs driven by a visual task as well as epileptogenic HFOs arising from the extraoccipital seizure focus. We identified spontaneous HFOs at ≥80 Hz with a mean duration of 330 ms intermittently emerging from the occipital cortex during interictal slow-wave sleep. The spectral frequency band of spontaneous occipital HFOs was similar to that of visually driven HFOs. Spontaneous occipital HFOs were spatially sparse and confined to smaller areas, whereas visually driven HFOs involved the larger areas including the more rostral sites. Neither spectral frequency band nor amplitude of spontaneous occipital HFOs significantly differed from those of epileptogenic HFOs. Spontaneous occipital HFOs were strongly locked to the phase of delta activity, but the strength of δ-phase coupling decayed from 1 to 3 Hz. Conversely, epileptogenic extraoccipital HFOs were locked to the phase of delta activity about equally in the range from 1 to 3 Hz. The occipital cortex spontaneously generates physiological HFOs which may stand out on electrocorticography traces as prominently as pathological HFOs arising from elsewhere; this observation should be taken into consideration during presurgical evaluation. Coupling of spontaneous delta and HFOs may increase the understanding of significance of δ-oscillations during slow-wave sleep. Further studies are warranted to determine whether δ-phase coupling distinguishes physiological from pathological HFOs or simply differs across anatomical locations.

γ-oscillations modulated by picture naming and word reading: intracranial recording in epileptic patients.

OBJECTIVE: We measured cortical gamma-oscillations in response to visual-language tasks consisting of picture naming and word reading in an effort to better understand human visual-language pathways. METHODS: We studied six patients with focal epilepsy who underwent extraoperative electrocorticography (ECoG) recording. Patients were asked to overtly name images presented sequentially in the picture naming task and to overtly read written words in the reading task. RESULTS: Both tasks commonly elicited gamma-augmentation (maximally at 80-100 Hz) on ECoG in the occipital, inferior-occipital-temporal and inferior-Rolandic areas, bilaterally. Picture naming, compared to reading task, elicited greater gamma-augmentation in portions of pre-motor areas as well as occipital and inferior-occipital-temporal areas, bilaterally. In contrast, word reading elicited greater gamma-augmentation in portions of bilateral occipital, left occipital-temporal and left superior-posterior-parietal areas. Gamma-attenuation was elicited by both tasks in portions of posterior cingulate and ventral premotor-prefrontal areas bilaterally. The number of letters in a presented word was positively correlated to the degree of gamma-augmentation in the medial occipital areas. CONCLUSIONS: Gamma-augmentation measured on ECoG identified cortical areas commonly and differentially involved in picture naming and reading tasks. Longer words may activate the primary visual cortex for the more peripheral field. SIGNIFICANCE: The present study increases our understanding of the visual-language pathways.

Somatosensory-related gamma-, beta- and alpha-augmentation precedes alpha- and beta-attenuation in humans.

OBJECTIVE: Several human studies have demonstrated that the amplitudes of cortical oscillations are altered by various sensorimotor and cognitive tasks. Event-related augmentation of gamma oscillations and attenuation of alpha and beta oscillations have been often used as surrogate markers of cortical activation elicited by tasks especially in presurgical identification of eloquent cortices. In the present study, we addressed a question whether somatosensory-related gamma augmentation 'precedes' or 'co-occurs with' somatosensory-related attenuation of alpha-beta oscillations. METHODS: We studied 10 patients who underwent intracranial electrocorticography for epilepsy surgery, and determined the temporal and spatial characteristics of median-nerve somatosensory-related amplitude changes at gamma- (30-100Hz), beta- (14-28Hz) and alpha-band (8-12Hz) oscillations. RESULTS: We found that somatosensory-related gamma augmentation involving the post- and pre-central gyri evolved into beta and alpha augmentation, which was subsequently followed by beta and alpha attenuation involving the post- and pre-central gyri. CONCLUSIONS: These observations support the hypothesis that somatosensory-related gamma augmentation but not alpha-beta attenuation represents the initial cortical processing for external somatosensory stimuli. Somatosensory-related alpha-beta attenuation appears to represent a temporally distinct stage of somatosensory processing. SIGNIFICANCE: The present study has increased our understanding of event-related gamma augmentation and alpha-beta attenuation seen on electrocorticography.

Superior temporal gyrus spectral abnormalities in schizophrenia.

Considerable evidence indicates early auditory stimulus processing abnormalities in schizophrenia, but the mechanisms are unclear. The present study examined oscillatory phenomena during a paired-click paradigm in the superior temporal gyrus (STG) as a possible core problem. The primary question addressed is whether first click and/or second click group differences in the time-domain evoked response in patients with schizophrenia are due to (1) group differences in the magnitude of poststimulus oscillatory activity, (2) group differences in poststimulus phase-locking, and/or (3) group differences in the magnitude of ongoing background oscillatory activity. Dense-array magnetoencephalography from 45 controls and 45 patients with schizophrenia produced left- and right-hemisphere STG 50- and 100-ms time-frequency evoked, phase-locking, and total power measures. Whereas first click 100-ms evoked theta and alpha abnormalities were observed bilaterally, evoked low beta-band differences were specific to the left hemisphere. Compared to controls, patients with schizophrenia showed more low-frequency phase variability, and the decreased 100-ms S1 evoked response observed in patients was best predicted by the STG phase-locking measure.

In vivo animation of auditory-language-induced gamma-oscillations in children with intractable focal epilepsy.

We determined if high-frequency gamma-oscillations (50- to 150-Hz) were induced by simple auditory communication over the language network areas in children with focal epilepsy. Four children (aged 7, 9, 10 and 16 years) with intractable left-hemispheric focal epilepsy underwent extraoperative electrocorticography (ECoG) as well as language mapping using neurostimulation and auditory-language-induced gamma-oscillations on ECoG. The audible communication was recorded concurrently and integrated with ECoG recording to allow for accurate time lock on ECoG analysis. In three children, who successfully completed the auditory-language task, high-frequency gamma-augmentation sequentially involved: i) the posterior superior temporal gyrus when listening to the question, ii) the posterior lateral temporal region and the posterior frontal region in the time interval between question completion and the patient's vocalization, and iii) the pre- and post-central gyri immediately preceding and during the patient's vocalization. The youngest child, with attention deficits, failed to cooperate during the auditory-language task, and high-frequency gamma-augmentation was noted only in the posterior superior temporal gyrus when audible questions were given. The size of language areas suggested by statistically significant high-frequency gamma-augmentation was larger than that defined by neurostimulation. The present method can provide in vivo imaging of electrophysiological activities over the language network areas during language processes. Further studies are warranted to determine whether recording of language-induced gamma-oscillations can supplement language mapping using neurostimulation in presurgical evaluation of children with focal epilepsy.

Novelty and target processing during an auditory novelty oddball: a simultaneous event-related potential and functional magnetic resonance imaging study.

Recent evidence suggests that both spatiotemporally distinct and overlapping brain regions are involved in bottom-up- and top-down-driven attentional processing. However, existing studies are based on a variety of different approaches, including electroencephalography (EEG) and functional magnetic resonance imaging (fMRI), raising the question of how EEG and fMRI findings in this field are related to each other. The present study aimed at disentangling common from specific regions underlying bottom-up novelty-processing and top-down target-processing. Simultaneous EEG and fMRI recordings were employed to investigate how fMRI-identified brain regions contribute to event-related potential (ERP) signatures of novelty- and target-processing. Fourteen subjects performed a modified novelty oddball task in which either rare tones or novel sounds served as targets in different blocks, allowing us to separate novelty-related from mere distractor-related effects. ERP signatures of novelty- and target-processing could be identified, confirming previous research based on recordings outside the scanner. fMRI analyses revealed that, despite considerable overlap of regions activated during novelty- and target-processing, bilateral superior temporal and right inferior frontal areas showed pronounced activation related to novelty-processing. fMRI-informed ERP dipole seeding was used to integrate both signals. The source modeling results further implicated temporal and inferior frontal sources in novelty-processing. Target-related fMRI activation on the other hand was confirmed in a network comprising distributed frontoparietal regions as well as bilateral caudate nucleus and cerebellum. Most regions identified by fMRI showed a contribution to target-related ERP signatures. This pattern of findings underscores the potential of simultaneous EEG/fMRI recordings for the spatiotemporal characterization of target- and novelty-processing.

Artifact correction and source analysis of early electroencephalographic responses evoked by transcranial magnetic stimulation over primary motor cortex.

Analyzing the brain responses to transcranial magnetic stimulation (TMS) using electroencephalography (EEG) is a promising method for the assessment of functional cortical connectivity and excitability of areas accessible to this stimulation. However, until now it has been difficult to analyze the EEG responses during the several tens of milliseconds immediately following the stimulus due to TMS-induced artifacts. In the present study we show that by combining a specially adapted recording system with software artifact correction it is possible to remove a major part of the artifact and analyze the cortical responses as early as 10 ms after TMS. We used this methodology to examine responses of left and right primary motor cortex (M1) to TMS at different intensities. Based on the artifact-corrected data we propose a model for the cortical activation following M1 stimulation. The model revealed the same basic response sequence for both hemispheres. A large part of the response could be accounted for by two sources: a source close to the stimulation site (peaking approximately 15 ms after the stimulus) and a midline frontal source ipsilateral to the stimulus (peaking approximately 25 ms). In addition the model suggests responses in ipsilateral temporo-parietal junction areas (approximately 35 ms) and ipsilateral (approximately 30 ms) and middle (approximately 50 ms) cerebellum. Statistical analysis revealed significant dependence on stimulation intensity for the ipsilateral midline frontal source. The methodology developed in the present study paves the way for the detailed study of early responses to TMS in a wide variety of brain areas.

Mental chronometry of working memory retrieval: a combined functional magnetic resonance imaging and event-related potentials approach.

We used the combination of functional magnetic resonance imaging and event-related potentials to decompose the processing stages (mental chronometry) of working memory retrieval. Our results reveal an early transient activation of inferotemporal cortex, which was accompanied by the onset of a sustained activation of posterior parietal cortex. We furthermore observed late transient responses in ventrolateral prefrontal cortex and late sustained activity in medial frontal and premotor areas. We propose that these neural signatures reflect the cognitive stages of task processing, perceptual evaluation (inferotemporal cortex), storage buffer operations (posterior parietal cortex), active retrieval (ventrolateral prefrontal cortex), and action selection (medial frontal and premotor cortex). This is also supported by their differential temporal contribution to specific subcomponents of the P300 cognitive potential.

EEG source analysis and fMRI reveal two electrical sources in the fronto-parietal operculum during subepidermal finger stimulation.

Using functional magnetic resonance imaging (fMRI) and electroencephalographic (EEG) source dipole analysis in 10 normal subjects, two electrical source dipoles in the contralateral fronto-parietal operculum were identified during repetitive painful subepidermal stimulation of the right index finger. The anterior source dipole peaking at 79 +/- 8 ms (mean +/- SD) was located in the frontal operculum, and oriented tangentially toward the cortical surface. The posterior source dipole peaking at 118 +/- 12 ms was located in the upper bank of the Sylvian fissure corresponding to the second somatosensory cortex (S2). The orientations of the posterior source dipoles displayed large variability, but differed significantly (P < 0.05) from the orientations of the anterior source dipoles. Electrical sources and fMRI clusters were also observed in ipsilateral fronto-parietal operculum. However, due to low signal-to-noise ratio of ipsilateral EEG sources in individual recordings, separation of sources into anterior and posterior clusters was not performed. Combined fMRI and source dipole EEG analysis of individual data suggests the presence of two distinct electrical sources in the fronto-parietal operculum participating in processing of somatosensory stimuli. The anterior region of the fronto-parietal operculum shows earlier peak activation than the posterior region.

Localizing P300 generators in visual target and distractor processing: a combined event-related potential and functional magnetic resonance imaging study.

Constraints from functional magnetic resonance imaging (fMRI) were used to identify the sources of the visual P300 event-related potential (ERP). Healthy subjects performed a visual three-stimulus oddball paradigm with a difficult discrimination task while fMRI and high-density ERP data were acquired in separate sessions. This paradigm allowed us to differentiate the P3b component of the P300, which has been implicated in the detection of rare events in general (target and distractor), from the P3a component, which is mainly evoked by distractor events. The fMRI-constrained source model explained >99% of the variance of the scalp ERP for both components. The P3b was mainly produced by parietal and inferior temporal areas, whereas frontal areas and the insula contributed mainly to the P3a. This source model reveals that both higher visual and supramodal association areas contribute to the visual P3b and that the P3a has a strong frontal contribution, which is compatible with its more anterior distribution on the scalp. The results point to the involvement of distinct attentional subsystems in target and distractor processing.

BESA source coherence: a new method to study cortical oscillatory coupling.

This paper introduces source coherence, a new method for the analysis of cortical coherence using noninvasive EEG and MEG data. Brain electrical source analysis (BESA) is applied to create a discrete multiple source model. This model is used as a source montage to transform the recorded data from sensor level into brain source space. This provides source waveforms of the modeled brain regions as a direct measure for their activities on a single trial basis. The source waveforms are transformed into time-frequency space using complex demodulation. Magnitude-squared coherence between the brain sources reveals oscillatory coupling between sources. This procedure allows one to separate the time-frequency content of different brain regions even if their activities severely overlap at the surface. Thus, source coherence overcomes problems of localization and interpretation that are inherent to coherence analysis at sensor level. The principle of source coherence is illustrated using an EEG recording of an error-related negativity as an example. In this experiment the subject performed a visuo-motor task. Source coherence analysis revealed dynamical linking between posterior and central areas within the gamma-band around the time of button press at a post-stimulus latency of 200-300 ms.

Source activity in the human secondary somatosensory cortex depends on the size of corpus callosum.

If corpus callosum (CC) mediates the activation of the secondary somatosensory area (SII) ipsilateral to the side of stimulation, then the peak latencies of the contra- and ipsilateral SII activity as well as the amplitude of the ipsilateral SII activity should correlate with the size of CC. Innocuous electrical stimuli of five different intensities were applied to the ventral surface of the right index finger in 15 right-handed men. EEG was recorded using 82 closely spaced electrodes. The size of CC and of seven callosal regions was measured from the mid-sagittal slice of a high-resolution anatomical MRI. The activation in the contralateral and ipsilateral SII was evaluated using spatio-temporal source analysis. At the strongest stimulus intensity, the size of the intermediate part of the callosal truncus correlated negatively with the interpeak latency of the sources in ipsi- and contralateral SII (r = -0.83, P < 0.01). Stepwise regression analysis showed that the large size of the intermediate truncus of CC was paralleled by a latency reduction of peak activity of the ipsilateral SII, whereas both contra- and ipsilateral peak latencies were positively correlated. The peak amplitude of the ipsilateral SII source correlated positively with the size of the intermediate truncus of CC, and with the peak amplitudes of sources in the primary somatosensory cortex (SI) and in the mesial frontal cortex. The results suggest that in right-handed neurologically normal men, the size of the intermediate callosal truncus contributes to the timing and amplitude of ipsilateral SII source activity.

Psychogenic sensory loss: magnetic source imaging reveals normal tactile evoked activity of the human primary and secondary somatosensory cortex.

We investigated the activation of the secondary somatosensory cortex (SII) in three patients with unilateral psychogenic sensory loss in the hand. Somatosensory evoked magnetic fields were recorded in response to tactile stimulation of the index finger of the affected and the unaffected hand. Brain sources were estimated by magnetic source imaging. In all subjects, responses in both the contralateral primary somatosensory (SI) and bilateral SII areas were normal as compared with a healthy control group, irrespective of the stimulated side. The results extend previous findings of normal evoked activity in SI.