Keiichiro Toma

Generators of movement-related cortical potentials: fMRI-constrained EEG dipole source analysis.

To clarify the precise location and timing of the mo tor cortical activation in voluntary movement, dipole source analysis integrating multiple constraints wa conducted for the movement-related cortical potentia (MRCP). Six healthy subjects performed single self paced extensions of the right index finger at about 15-intervals during EEG and event-related fMRI acquisi tions. EEG was recorded from 58 scalp electrodes, and fMRI of the entire brain was obtained every 2.6 s. Coordinates of the two methods were coregistered us ing anatomical landmarks. During dipole source mod eling, a realistic three-layer head model was used as a volume conductor. To identify the number of uncorre lated source s in the MRCP, principal component (PC analysis was performed, which was consistent with the existence of six sources in the left (Lt SM1) and right (Rt SMI) sensorimotor and medial frontocentral (MFC) areas. After dipoles were seeded at the acti vated spots revealed by fMRI, dipole orientations were fixed based on the interpretation of the topography of distribution of the PC. The strength of the six dipoles (three dipoles in Lt SMI, two in Rt SMI, and one in MFC) was then computed over time. Within the bilat eral SM1, activation of the precentral gyrus occurs bilaterally with similar strength from -1.2 s, followed by that of the precentral bank from -0.5 s with con tralateral preponderance. Subsequently, the postcen tral bank becomes active only on the contralateral side at 0.1 s after movement. Activation of the MFC shows timing similar to that of the bilateral precentral gyri These deduced patterns of activation are consis tent with previous studies of electrocorticography in humans.

Short-term high-frequency transcutaneous electrical nerve stimulation decreases human motor cortex excitability.

Several previous studies have shown that periods of changed sensory input can have after effects on the excitability of the corticospinal system. Here we test whether the parameters of peripheral stimulation conventionally used to treat pain with transcutaneous electrical nerve stimulation (TENS: 90 Hz) also have modulatory effects on the motor system. We measured the amplitude of motor evoked potentials (MEPs) elicited by the focal transcranial magnetic stimulation in the right abductor pollicis brevis and first dorsal interosseous muscles before and after 30 min TENS over the right thenar eminence. In addition, we evaluated tactile and 2-point discrimination thresholds at the same site. TENS transiently reduced MEPs and increased sensory thresholds. This suggests that short-term TENS might have an inhibitory effect on both the sensory and motor systems.

Implicit and explicit processing of kanji and kana words and non-words studied with fMRI

Using functional magnetic resonance imaging (fMRI), we investigated the implicit language processing of kanji and kana words (i.e., hiragana transcriptions of normally written kanji words) and non-words. Twelve right-handed native Japanese speakers performed size judgments for character stimuli (implicit language task for linguistic stimuli), size judgments for scrambled-character stimuli (implicit language task for non-linguistic stimuli), and lexical decisions (explicit language task). The size judgments for scrambled-kanji stimuli and scrambled-kana stimuli produced activations on the bilateral lingual gyri (BA 18), the bilateral occipitotemporal regions (BA 19/37), and the bilateral superior and inferior parietal cortices (BA 7/40). Interestingly, besides these areas, activations of the left inferior frontal region (Brocas area, BA 44/45) and the left posterior inferior temporal cortex (PITC, BA 37), which have been considered as language areas, were additionally activated during size judgment for kanji character stimuli. Size judgment for kana character stimuli also activated Brocas area, the left PITC, and the left supramarginal gyrus (SMG, BA 40). The activations of these language areas were replicated in the lexical decisions for both kanji and kana. These findings suggest that language processing of both kanji and kana scripts is obligatory to literate Japanese subjects. Moreover, comparison between the scrambled kanji and the scrambled kana showed no activation in the language areas, while greater activation in the bilateral fusiform gyri (left-side predominant) was found in kanji vs. kana comparison during the size judgment and the lexical decision. Kana minus kanji activated the left SMG during the size judgment, and Brocas area and the left middle/superior temporal junction during the lexical decision. These results probably reflect that in implicit or explicit reading of kanji words and kana words (i.e., hiragana transcriptions of kanji words), although using largely overlapping cortical regions, there are still some differences. Kanji reading may involve more heavily visual orthographic retrieval and lexical-semantic system through the ventral route, while kana transcriptions of kanji words require phonological recoding to gain semantic access through the dorsal route.

Second somatosensory area (SII) plays a significant role in selective somatosensory attention

In order to explore human cortical areas involved in active attention toward a somatosensory modality, somatosensory evoked cortical magnetic fields were recorded in ten healthy adults with a 122-channel whole-head magnetometer while the subjects performed the selective attention task. Two kinds of stimulus modality, somatosensory and auditory, were presented independently in the same session. For the somatosensory modality, a randomized sequence of strong (P=0.45) and weak (P=0.05) electric stimuli was delivered to the right median nerve at the wrist. For the auditory modality, a randomized sequence of 900-Hz (P=0.45) and 950-Hz (P=0.05) tones was delivered to both ears. Subjects were requested to pay attention to the specified stimulus modality (either somatosensory or auditory) and to count the number of rare stimuli of the attended modality (weak stimuli in the somatosensory or 950-Hz tone in the auditory modality). A total of 12 sessions were performed for each subject, among which the order of attended modality was changed alternately and counterbalanced among subjects. In the data analysis, somatosensory evoked fields for frequent stimuli (strong electric stimuli) were compared between the two conditions; attend somatosensory condition (ATS) and attend auditory condition (non-attend somatosensory condition; NATS). In six out of the ten subjects, somatosensory evoked fields showed attention-related change. The magnitude of the estimated generator source in SII, but not in SI, significantly increased from NATS to ATS while keeping the same locations. Moreover, a simulation study using the estimated sources in SII in NATS supported the enhancement of the activity in the SII rather than participation of additional sources in the selective attention task. These results suggest that the SII plays a main role in selective somatosensory attention.

Discrimination of Exner's area and the frontal eye field in humans--functional magnetic resonance imaging during language and saccade tasks.

In the left frontal lobe, Exners area (EXA), which is responsible for writing and reading, is located close to the frontal eye field (FEF), which is responsible for eye movements. To discriminate EXA from FEF anatomically and functionally, functional magnetic resonance imaging was conducted in 12 healthy volunteers. The saccadic eye movement experiment activated a region defined as the FEF, whereas three language experiments that included translation between grapheme and phoneme activated another region defined as EXA. EXA was found to be located only 1.5 cm apart from the FEF in the Talairach brain template. By conducting the saccade and language experiments in the same individuals, this study was able to successfully separate EXA from FEF.

Activation of the precuneus is related to reduced reaction time in serial reaction time tasks

Multiple brain areas are activated during serial reaction time (RT) tasks (SRTTs), but the part of the brain that facilitates reductions in RT remains unclear. The present study attempted to determine the brain region contributing most to improved RTs during explicit SRTTs. Subjects comprised 18 healthy volunteers who were instructed to press one of four buttons corresponding to visual stimuli as quickly as possible and with minimal errors during functional MRI. Stimuli were presented either in random order (control condition) or in a repeated six-item sequence (learning condition). Conventional analysis contrasting learning and control conditions revealed activation in the prefrontal-parietal area, which shifted to motor area. Subjects with high RT reduction showed more prominent activation in the precuneus than subjects with low RT reduction. Intra-subject correlation analysis revealed that time course of precuneus activation was unrelated to time-course of RT reduction. However, inter-subject correlation analysis revealed that RT changes correlate only with precuneus activation, meaning that subjects showing more prominent RT reduction revealed more prominent activation of the precuneus, which is known to play critical roles in controlling finger movements with reference to buffered memory.

A functional magnetic resonance imaging study of internal modulation of an external visual cue for motor execution

The strategy to perform a task differs according to how a cue is interpreted. In order to investigate the basic mechanisms of temporal regulation in the higher motor areas, the interaction between two different types of internal modulations of an external visual cue was evaluated using functional magnetic resonance imaging (fMRI). An opposing finger movement task guided by dot prompting was employed. In the intermittent tapping experiment, two taps per second and a rest for one second were alternatively repeated in the task blocks. In the constant tapping experiments, the volunteers performed finger movements at 0.5, 1 or 2 Hz. The activation in the primary sensory motor area correlated with the amount of movement. Activities in the supplementary motor area, left dorsal pre-motor area, left superior parietal lobule and right cerebellum depended on the demand for internal modulation. Activation in these areas was maximum for the intermittent task which was a combination of two different internal modulations, and minimum for the 1 Hz movement that did not require internal modulation. It was suggested that these four areas are directly involved in the generation of a complex movement sequence driven by a visual cue, and they are organized for performance. The translation of external pacing and initiation for self-pacing may share the same neuronal basis. Activation in the left supramarginal gyrus, bilateral frontal operticula and basal ganglia did not depend on the combination of the two internal modulations.

Subregions of human MT complex revealed by comparative MEG and direct electrocorticographic recordings

OBJECTIVE To locate the visual motion complex (MT+) and study its response properties in an epilepsy surgery patient. METHODS A 17-year-old epilepsy patient underwent invasive monitoring with subdural electrodes in the right temporo-parieto-occipital area. MT+ was investigated by cortical electric stimulation and by epicortical visual evoked potentials time-locked to motion onset of sinusoidal gratings (motion VEP). Motion-related visual evoked magnetic field (motion VEF) was also recorded before the electrode implantation to complement the invasive recording. RESULTS Motion VEPs revealed two subregions within MT+, generating early and late potentials respectively. The early activity with a peak around 130 ms was localized at a single electrode situated immediately caudal to the initial portion of the ascending limb of the superior temporal sulcus (AL-STS). The late activity, peaking at 242-274 ms, was located ventro-rostrally over three electrodes. Among the four electrodes with motion VEPs, cortical stimulation at the most caudal pair elicited motion-in-depth perception involving the whole visual field. In addition to two subregions revealed on the gyral crown, magnetoencephalography (MEG) demonstrated another subregion with a late motion VEF in AL-STS immediately rostral to the electrode with the early motion VEP. CONCLUSIONS In combination with MEG recording, the present invasive exploration demonstrated human MT+ in a focal area of the temporo-parieto-occipital junction and delineated possible three subregions as indicated by the different latencies and distributions of the motion VEP/VEFs. SIGNIFICANCE Comparative MEG and direct electrocorticographic recordings delineated possible subregions within the human MT complex.

Generators of the Movement-Related Cortical Potentials and Dipole Source Analysis

The precise location and timing of cortical activation in voluntary movements have been major issues in motor control physiology. For example, during movement preparation, whether the supplementary motor area (SMA) is activated in a sequential (Deecke and Kornhuber, 1978; Orgogozo and Lasen, 1979; Deecke, 1987; Deecke and Lang, 1996; Deecke et al., 1999) or parallel (Hyland et al., 1989; Ikeda et al., 1992) manner has been intensively debated with regard to the primary motor area (M1). During movement execution, contribution of the gyrus and sulcus part of motor cortex has also been an important issue (Strick and Preston, 1982a, Strick and Preston, 1982b; Kawashima et al., 1995; Geyer et al., 1996). Intracranial recordings with subdural (Neshige et al., 1988; Ikeda et al., 1992) as well as depth (Rektor et al., 1994) electrodes are useful approaches to gain information about the precise cortical regions and time course of the activation. However, these techniques invasively record pathological brains, and the limitations of electrode placement over distributed cortical regions prevent simultaneous collection of data from various brain areas. In addition, subdural recording from an electrode placed over the gyrus provides little information about activity from the sulcus. Recent development and improvement of electromagnetic or neuroimaging measurements enable us to investigate human brain function noninvasively. At the present time, however, no non-invasive methodology with sufficient temporal and spatial resolution has been developed. Electromagnetic measurements such as the electroencephalogram (EEG) and magnetoencephalogram (MEG) provide information about cortical activation with millisecond resolution, but with limited spatial localization. By contrast, neuroimaging techniques such as positron emission tomography (PET) (Hersovitch, 1994) and functional magnetic resonance imaging (fMRI) (Friston et al., 1995; et al., 1998) yield good spatial information with millimeter resolution, although they are devoid of good timing information.

The role of the human supplementary motor area in reactive motor operation.

The role of the supplementary motor area (SMA) in reactive motor operation was investigated with functional magnetic resonance imaging in 13 normal subjects. A visual cue was presented at a regular (1 Hz) or irregular (mean, 1 Hz) rate, and the subject pressed a button with the right index finger in a predictive or reactive manner. Brain regions associated with reactive movement were detected by comparing reactive with predictive movement tasks, and those with irregular movement by comparing irregular and regular cueing tasks. During regular cueing, the SMA showed greater activation for reactive than predictive movement. However, the SMA was equally activated between regular and irregular cueing once the subject reacted to the cue. The SMA is likely involved in reactive adjustment of movement to the external cue.