Development of a generative model of magnetoencephalography noise that enables brain signal extraction from single-epoch data.

We presented a method of rejecting sensor-specific and environmental noise during magnetoencephalography (MEG) measurement that enables the extraction of brain signals from single-epoch data. The method assumes a parametric generative model of MEG data. The model's optimal parameters were determined from single-epoch data, and noise reduction was performed by the decomposition of data within the optimal model. We confirmed our method's validity through multiple experiments. Moreover, we compared our method's performance with that of several previous noise-reduction methods. Finally, we confirmed that the proposed method followed by spatial filtering reduced noise more efficiently.

Human neural responses involved in spatial pooling of locally ambiguous motion signals.

Early visual motion signals are local and one-dimensional (1-D). For specification of global two-dimensional (2-D) motion vectors, the visual system should appropriately integrate these signals across orientation and space. Previous neurophysiological studies have suggested that this integration process consists of two computational steps (estimation of local 2-D motion vectors, followed by their spatial pooling), both being identified in the area MT. Psychophysical findings, however, suggest that under certain stimulus conditions, the human visual system can also compute mathematically correct global motion vectors from direct pooling of spatially distributed 1-D motion signals. To study the neural mechanisms responsible for this novel 1-D motion pooling, we conducted human magnetoencephalography (MEG) and functional MRI experiments using a global motion stimulus comprising multiple moving Gabors (global-Gabor motion). In the first experiment, we measured MEG and blood oxygen level-dependent responses while changing motion coherence of global-Gabor motion. In the second experiment, we investigated cortical responses correlated with direction-selective adaptation to the global 2-D motion, not to local 1-D motions. We found that human MT complex (hMT+) responses show both coherence dependency and direction selectivity to global motion based on 1-D pooling. The results provide the first evidence that hMT+ is the locus of 1-D motion pooling, as well as that of conventional 2-D motion pooling.

Visual mismatch response evoked by a perceptually indistinguishable oddball.

Mismatch field (MMF) is an early magnetoencephalographic response evoked by deviant stimuli within a sequence of standard stimuli. Although auditory MMF is reported to be an automatic response, the automaticity of visual MMF has not been clearly demonstrated, partly because of the difficulty in designing an ignore condition. Our modified oddball paradigm had a masking stimulus inserted between briefly presented standard and deviant stimuli (vertical gratings with different spatial frequencies). Perceptual discrimination between masked standard and deviant stimuli was difficult, but the early magnetoencephalographic response for the deviant was significantly larger than that for the standard, when the former had a higher spatial frequency than the latter. Our findings strongly support the hypothesis that visual MMF is evoked automatically.

Neural processes for intentional control of perceptual switching: a magnetoencephalography study.

This article reports an interesting link between the psychophysical property of intentional control of perceptual switching and the underlying neural activities. First, we revealed that the timing of perceptual switching for a dynamical dot quartet can be controlled by the observers' intention, without eye movement. However, there is a clear limitation to this control, such that each animation frame of the stimulus must be presented for a sufficiently long time length; in other words, the frequency of the stimulus alternation must be sufficiently slow for the control. The typical stimulus onset asynchrony for a 50% level of success was about 275 ms for an average of 10 observers. On the basis of psychophysical property, we designed three experiments for investigating the neural process with a magnetoencephalography. They revealed that: (1) a peak component occurring about 300 ms after a reversal was stronger when the direction of perceived motion was switched intentionally than when it was not switched, and (2) neural components about 30-40 ms and 240-250 ms after the reversal of the stimulus animation were stronger when perception was altered intentionally than when it was switched unintentionally. The 300 ms component is consistent with a previous study about passive perceptual switching (Struber and Herrmann [ 2002]: Cogn Brain Res 14:370-382), but the intentional effect was seemed to be a different component from the well-known P300 component.

Dissociable neural correlates of reorienting within versus across visual hemifields.

Neural correlates of reorienting across visual hemifields have been extensively studied, however, those of reorienting within hemifields and if there are any differences remain unclear. Here, we performed a functional magnetic resonance imaging study to identify neural correlates of reorienting within and across hemifields using a variant of the cueing paradigm. Behavioral results showed that reorienting across hemifields showed significant validity effect, but reorienting within hemifields did not. Functional magnetic resonance imaging data revealed dissociable activations in the right posterior parietal region between reorienting within and across hemifields. The present results suggest that reorienting within hemifields differs from the 'classical reorienting' to some extent, whereas reorienting across hemifields does not.

Close similarity between spatiotemporal frequency tunings of human cortical responses and involuntary manual following responses to visual motion.

Human brain uses visual motion inputs not only for generating subjective sensation of motion but also for directly guiding involuntary actions. For instance, during arm reaching, a large-field visual motion is quickly and involuntarily transformed into a manual response in the direction of visual motion (manual following response, MFR). Previous attempts to correlate motion-evoked cortical activities, revealed by brain imaging techniques, with conscious motion perception have resulted only in partial success. In contrast, here we show a surprising degree of similarity between the MFR and the population neural activity measured by magnetoencephalography (MEG). We measured the MFR and MEG induced by the same motion onset of a large-field sinusoidal drifting grating with changing the spatiotemporal frequency of the grating. The initial transient phase of these two responses had very similar spatiotemporal tunings. Specifically, both the MEG and MFR amplitudes increased as the spatial frequency was decreased to, at most, 0.05 c/deg, or as the temporal frequency was increased to, at least, 10 Hz. We also found in peak latency a quantitative agreement (approximately 100-150 ms) and correlated changes against spatiotemporal frequency changes between MEG and MFR. In comparison with these two responses, conscious visual motion detection is known to be most sensitive (i.e., have the lowest detection threshold) at higher spatial frequencies and have longer and more variable response latencies. Our results suggest a close relationship between the properties of involuntary motor responses and motion-evoked cortical activity as reflected by the MEG.

Alpha band amplification during illusory jitter perception.

Synchronization is thought to have a role in linking disparate components into neural assemblies. However, the particular frequency of the synchronization is generally considered to be incidental to its functional role. Here we report a link between enhanced alpha activations and an illusory jitter of the same frequency. We measured perceived jitter rates and the magnetoencephalography during presentations of a stimulus wherein red squares and superimposed vertical green bars moved together across a black background. The green bars were either darker, equiluminant with, or brighter than the red squares. We established that the illusory jitter rate, robustly seen only in the equiluminant condition, was approximately 10 Hz. Crucially, neural oscillations around 10 Hz were enhanced in this condition. Surprisingly, approximately 10 Hz oscillations were also enhanced during illusory jitter perception relative to a moving stimulus that contained physical 10 Hz jitter. This suggests that the enhanced synchronization is associated with illusory jitter generation rather than with jitter perception. Since the stimulus eliciting illusory jitter moves smoothly and rigidly, both the percept and enhanced neural synchrony must be generated within the visual system. Our data therefore indicate a match between the dynamics of synchronous neural activity and the dynamics of a sensory experience offering the intriguing possibility of a common cause.

Predictive and sensory integration begins at an early stage of visual processing.

Brain activity was measured by magnetoencephalography to investigate the spatiotemporal stage of visual processing at which predictive and sensory integration begins. We examined the consequences of a visual mismatch between preliminary prediction and incoming stimulus. Following auditory cues (1000- and 1250-Hz tones) for prediction, congruent and incongruent images, pictures of two musical keys, were presented to volunteers. When they predicted visual inputs on the basis of preceding auditory cues, we detected a mismatch signal for predictive-sensory incongruities in the striate and extrastriate areas for 100-200 ms after image presentation. As this signal reflects a compatibility analysis, we propose that the integration process begins in these areas approximately 100 ms after image presentation.

Direct reconstruction algorithm of current dipoles for vector magnetoencephalography and electroencephalography.

This paper presents a novel algorithm to reconstruct parameters of a sufficient number of current dipoles that describe data (equivalent current dipoles, ECDs, hereafter) from radial/vector magnetoencephalography (MEG) with and without electroencephalography (EEG). We assume a three-compartment head model and arbitrary surfaces on which the MEG sensors and EEG electrodes are placed. Via the multipole expansion of the magnetic field, we obtain algebraic equations relating the dipole parameters to the vector MEG/EEG data. By solving them directly, without providing initial parameter guesses and computing forward solutions iteratively, the dipole positions and moments projected onto the xy-plane (equatorial plane) are reconstructed from a single time shot of the data. In addition, when the head layers and the sensor surfaces are spherically symmetric, we show that the required data reduce to radial MEG only. This clarifies the advantage of vector MEG/EEG measurements and algorithms for a generally-shaped head and sensor surfaces. In the numerical simulations, the centroids of the patch sources are well localized using vector/radial MEG measured on the upper hemisphere. By assuming the model order to be larger than the actual dipole number, the resultant spurious dipole is shown to have a much smaller strength magnetic moment (about 0.05 times smaller when the SNR = 16 dB), so that the number of ECDs is reasonably estimated. We consider that our direct method with greatly reduced computational cost can also be used to provide a good initial guess for conventional dipolar/multipolar fitting algorithms.

Functional modulation of power-law distribution in visual perception.

Neuronal activities have recently been reported to exhibit power-law scaling behavior. However, it has not been demonstrated that the power-law component can play an important role in human perceptual functions. Here, we demonstrate that the power spectrum of magnetoencephalograph recordings of brain activity varies in coordination with perception of subthreshold visual stimuli. We observed that perceptual performance could be better explained by modulation of the power-law component than by modulation of the peak power in particular narrow frequency ranges. The results suggest that the brain operates in a state of self-organized criticality, modulating the power spectral exponent of its activity to optimize its internal state for response to external stimuli.

A comparison of stimulus synchronous activity in the primary motor cortices of athletes and non-athletes.

In this study, we measured primary motor cortex (MI) activity during a reaction time task to examine the appearance of MI activity that synchronized with the stimulus presentation (stimulus synchronous MI activity, SSMA). Because brain activity was expected to be enhanced by the repetitive/extensive activation, we hypothesized that the SSMA would be more clearly observable in athletes who were trained to perform reactive movements than in non-athletes. MI activity was measured in ten athletes and ten non-athletes by magnetoencephalography. The tasks were a simple reaction task and a Go/Nogo reaction task in which the subjects were asked to abduct their right index fingers in response to a visual stimulus. The Go/Nogo reaction time task was adopted to confirm the presence of the SSMA, because the MI activity in response to a Nogo stimulus did not overlap with the MI activity that was synchronous with the execution of the movement. The results show that the SSMA was clearly apparent in the athlete group (9/10). In the non-athlete group, however, only three subjects showed the SSMA (3/10). Moreover, the MI activity of the athletes tended to be larger than that of the non-athletes, even though the athletes did not specifically practice these index finger movements during their daily training. We concluded that long-term physical training promotes MI activity and the effects of reactive task repetition were more clearly apparent in the MI activity of the athletes.

Estimation of the timing of human visual perception from magnetoencephalography.

To explore the timing and the underlying neural dynamics of visual perception, we analyzed the relationship between the manual reaction time (RT) to the onset of a visual stimulus and the time course of the evoked neural response simultaneously measured by magnetoencephalography (MEG). The visual stimuli were a transition from incoherent to coherent motion of random dots and an onset of a chromatic grating from a uniform field, which evoke neural responses in different cortical sites. For both stimuli, changes in median RT with changing stimulus strength (motion coherence or chromatic contrast) were accurately predicted, with a stimulus-independent postdetection delay, from the time that the temporally integrated MEG response crossed a threshold (integrator model). In comparison, the prediction of RT was less accurate from the peak MEG latency, or from the time that the nonintegrated MEG response crossed a threshold (level detector model). The integrator model could also account for, at least partially, intertrial changes in RT or in perception (hit/miss) to identical stimuli. Although we examined MEG-RT relationships mainly for data averaged over trials, the integrator model could show some correlations even for single-trial data. The model predictions deteriorated when only early visual responses presumably originating from the striate cortex were used as the input to the integrator model. Our results suggest that the perceptions for visual stimulus appearances are established in extrastriate areas [around MT (middle temporal visual area) for motion and around V4 (fourth visual area) for color] approximately 150-200 ms before subjects manually react to the stimulus.

MEG responses correlated with the visual perception of velocity change.

Magnetoencephalography (MEG) was used to find neural activities, in the human brain, involved in perception of velocity changes in visual motion. We recorded MEG responses evoked by the stimuli whose velocity increased by 40% or 80% of baseline velocities of 1.0, 2.0, 3.0, and 4.0 deg/s. The velocity increment threshold and the manual reaction time (RT) were also measured under similar stimulus conditions. To manipulate observer's sensitivity to velocity increments, the MEG responses and the psychophysical performances were measured after adaptation to motion in one direction (adapted condition) or alternating directions (control condition). MEG responses evoked by velocity increments peaked at 200-290 ms (M1), and the M1 amplitudes, especially those obtained for 40% increments, were correlated with the sensitivities, which are the reciprocal of velocity increment thresholds. Furthermore, motion adaptation enhanced sensitivity to velocity increments and increased the M1 amplitudes. These results suggest a close correlation between the perceptual velocity increment and the evoked MEG response. In other words, the results suggest that velocity increments are detectable when there is a constant increment in magnetic neural response. As for latencies, nearly constant value of M1 latency did not quantitatively match a large decrease in manual RT with the increase in the baseline velocity. Motion adaptation reduced neither the peak MEG latency nor the manual RT.

Modulation of early auditory processing by visually based sound prediction.

Brain activity was measured by magnetoencephalography (MEG) to investigate whether the early auditory system can detect changes in audio-visual patterns when the visual part is presented earlier. We hypothesized that a template underlying the mismatch field (MMF) phenomenon, which is usually formed by past sound regularities, is also used in visually based sound prediction. Activity similar to the MMF may be elicited by comparing an incoming sound with the template. The stimulus was modeled after a keyboard: an animation in which one of two keys was depressed was accompanied by either a lower or higher tone. Congruent audio-visual pairs were designed to be frequent and incongruent pairs to be infrequent. Subjects were instructed to predict an incoming sound based on key movement in two sets of trials (prediction condition), whereas they were instructed not to do so in the other two sets (non-prediction condition). For each condition, the movement took 50 ms in one set (Delta = 50 ms) and 300 ms in the other (Delta = 300 ms) to reach the bottom, at which time a tone was delivered. As a result, only under the prediction condition with Delta = 300 ms was additional activity for incongruent pairs observed bilaterally in the supratemporal area within 100-200 ms of the auditory stimulus onset; this activity had spatio-temporal properties similar to those of MMF. We concluded that a template is created by the visually based sound prediction only after the visual discriminative and sound prediction processes have already been performed.

MEG recording from the human ventro-occipital cortex in response to isoluminant color stimulation.

In contrast to PET and fMRI studies, color-selective responses from the ventro-occipital area have rarely been reported in MEG studies. We tried to minimize the stimulation to all areas in the visual system except the color-processing ones by using a color space based on psychophysical and physiological knowledge in order to maximize the signal-to-noise ratio for MEG responses from the ventro-occipital area. MEG obtained from long intermittent reversals (2.0-3.5 s) of isoluminant chromatic gratings showed two major peaks at the latencies of approximately 100 and 150 ms. The estimated location of the equivalent-current dipole for response at 100-ms latency was in the calcarine sulcus and that of the dipole for the response at 150 ms was in the collateral sulcus in the ventro-occipital area. The response around 150 ms was uniquely observed in MEG elicited by chromatic reversals. The average of lags between MEG responses from the calcarine sulcus and ventro-occipital area was 43 ms, which suggests sequential processing of color information across the visual cortices.

Direction-specific adaptation of magnetic responses to motion onset.

We investigated the direction-specificity of motion adaptation, by recording magnetic responses evoked by motion onsets under both adapted and control conditions. The inter-stimulus interval was equated between the conditions to precisely evaluate the effect of motion adaptation itself. The onset stimuli at 1.5, 3.0 or 6.0 deg/s moved in the same direction or in the opposite direction to an adaptation stimulus at 3.0 deg/s. The perceived velocity of each test stimulus was measured in separate sessions. The most prominent peak (M2) of evoked responses appeared around 200-300 ms after motion onsets, and the dipoles were mainly estimated in the temporo-occipital area. Adaptation largely affected both perceived velocities and the M2 amplitudes. The M2 amplitudes were decreased by adaptation for both directions of test stimuli, and the decreases were significantly larger for the test stimuli in the adapted direction (49-63% of control condition) than for the test stimuli in the opposite direction (17-27% of control condition). The present study, for the first time, found that magnetic responses evoked by motion onsets reflect the activities of neurons that have direction-specificity.

The effect of 1/f fluctuation in inter-stimulus intervals on auditory evoked mismatch field.

This study focused on the effect of regularity of environmental stimuli on the informational order extracting function of human brain. The regularity of environmental stimuli can be described with the exponent n of the fluctuation 1/f(n). We studied the effect of the exponent of the fluctuation in the inter-stimulus interval (ISI) on the elicitation of auditory evoked mismatch fields (MMF) with two sounds with alternating frequency. ISI times were given by three types of fluctuation, 1/f(0), 1/f(1), 1/f(2), and with a fixed interval (1/f(infinity)). The root mean square (RMS) value of the MMF increased significantly (F(3/9)=4.95, p=0.027) with increases in the exponent of the fluctuation. Increments in the regularity of the fluctuation provoked enhancement of the MMF, which reflected the production of a memory trace, based on the anticipation of the stimulus timing. The gradient of the curve, indicating the ratio of increments between the MMF and the exponent of fluctuation, can express a subject's capability to extract regularity from fluctuating stimuli.

Bilateral cerebral activity for unilateral foot movement revealed by whole-head magnetoencephalography.

The primary motor cortices controlling foot movement are located on opposite sides of the longitudinal fissure. As a separation of closely located activity sources is not successful, the possibility of bilateral activation for lower limb movement remains undetermined. We therefore examined cerebral activity during unilateral foot movement to investigate the possibility of bilateral activation of primary foot motor cortices. Self-paced foot movement and finger movement (for comparison) were performed on ten subjects. Brain magnetic fields were recorded using a 64-channel whole-cortex magnetoencephalography (MEG) system. Brain activities were identified using 1- to 3-dipole models. Results evaluating finger movement were similar to previous reports. Equivalent current dipoles (ECDs) for foot movements were estimated in the primary foot motor and sensory regions. Sensory activity was always localized to the contralateral hemisphere. Motor activity was estimated by one ECD, but the laterality differed between subjects. Additional activity was discovered together with the primary motor activity, localized around the precentral sulcus. In contrast to consistent results of primary sensory activity, the variation of laterality of the foot primary motor ECD can be explained with a cancellation model, in which the magnetic fields generated from two closely spaced ECDs overlap to cancel each other out. Consequently, activation of the primary foot motor cortices was determined to be bilateral. Furthermore, it was estimated that additional activity may occur in the premotor area. This work suggests not only the bilateral activation of the primary foot motor cortices but also the possibility of a contribution of the premotor area.

A delayed class of BOLD waveforms associated with spreading depression in the feline cerebral cortex can be detected and characterised using independent component analysis (ICA).

An application of independent component analysis to blood oxygenation level- dependent MRI (BOLD-MRI) results was used to detect cerebrovascular changes that followed the initiation of cortical spreading depression (CSD) in feline brain. The cortical images were obtained from a horizontal plane at 28 s intervals before, and for 1.4-1.75 h after, KCl dissolved in agar (KCl/agar) had been directly applied to the left suprasylvian gyrus of 13 anesthetized cats for 10 min. It successfully resolved, for the first time, a novel class of prolonged, and delayed, biphasic BOLD waveforms. These were larger in amplitude ( approximately 20%), longer lasting and more delayed in onset (13-33 min) than the brief propagating (90 s) BOLD increase ( approximately 4%) already known to be associated with CSD on earlier occasions. Furthermore, such changes occurred in localized regions on the hemisphere ipsilateral to the site of stimulus application in 4 out of 5 control subjects rather than themselves generating propagating waves. Finally, the biphasic waveforms were consistently abolished in the 4 experimental animals studied following the i.v. administration of sumatriptan (0.3 mg kg(-1)), an antimigraine 5-HT(1B/1D) agonist, 15 min before the application of the transient stimulus. They were abolished in 2 out of 4 animals following the intraperitoneal (i.p.) administration of SB-220453 (tonabersat: 10 mg kg(-1), 90 min before stimulus application), a novel anticonvulsant that has recently been reported to inhibit CSD. ICA has thus been successful in detecting a novel localized, as opposed to propagating, signal of potential physiological significance hidden in complex BOLD- MRI data, whose sensitivity to sumatriptan may relate it to the cerebrovascular changes reported in the headache phase of migraine.

Magnetoencephalographic study of speed-dependent responses in apparent motion.

OBJECTIVES: There have been only few studies of visually-evoked cortical responses to apparent motion as a function of stimulus speed. Most earlier findings on evoked peak magnitudes and latencies, utilizing various types of smooth and apparent motion stimuli, have demonstrated that greater spatial separation/speed resulted in enhanced peak magnitudes, decreasing onset latencies in individual extrastriate neurons and in shorter motor reaction times in subjects. However, some reports using partial-coverage magnetoencephalography stated that increasing the stimulus displacement actually triggered a substantial reduction of the evoked main peak latency while the magnitude showed no clear change. METHODS: To resolve the issue of the dependency of evoked responses on stimulus speed in apparent motion, we presented moving bar stimuli to 6 subjects at velocities within a 100 fold range and investigated the ensuing evoked visual cortical activity using a whole-cortex magnetoencephalograph. The magnitude and the latency of the first major evoked peak M1 was measured and compared for 6 discrete bar-stimuli displacements in all subjects. RESULTS: Our results showed clearly that the M1 peak response magnitudes increased in a nonlinear way with higher apparent speeds (larger displacements), in compliance with the logarithmic Fechner law. We observed also that the fluctuations of the mean evoked M1 peak latency (140+/-10.6 ms) did not reach significance over the tested range of stimulus velocities. CONCLUSIONS: These findings probably reflect global motion processing mechanisms which rely on nonlinear speed-dependent feedback connectivity between striate and extrastriate visual cortex areas.