Trajectories of alpha rhythm dipoles shifting over the human brain cortex

Dynamic study of 3D localization of the equivalent current dipoles (ECD)--sources of the EEG alpha rhythm in the human brain was performed in seven subjects with closed eyes using a one-dipole model. An exact localization of ECDs was obtained by combination of EEG and MRI mapping that allowed tracing of ECD shifts over the cortex with 4 ms step. Our data confirmed localization of these ECDs mainly in the occipital cortex and revealed their successive shift over this area during generation of each alpha-wave. Typical trajectories of these shifts were revealed and quantitatively compared by the hierarchical cluster analysis. The data obtained directly proved periodical rhythmic alpha-wave spreading process in the human visual cortex and an external control of this process. The data are discussed in terms of the "scanning hypothesis" (Pitts W., McCulloch W.H. Bull. Math. Biophys. 1947. V. 9. P. 127) which predicted a certain functional meaning of the alpha activity for cortical processing of sensory information in the human brain.     

Neuromagnetic evidence that the P100 component of the pattern reversal visual evoked response originates in the bottom of the calcarine fissure

Visual evoked magnetic fields due to pattern reversal stimuli were measured in 5 normal subjects using a helmet-shaped 66 channel magnetoencephalography system linked to MRI. The magnetic topography of the prominent 100 ms response (P100m) evoked by full-field visual showed a double-dipole pattern in the occipital areas of all subjects. Right or left half-field stimuli and upper or lower quadrant-field stimuli evoked a single-dipole pattern in the contralateral occipital area. The P100m sources were when localized using a current dipole model and superimposed on MRI images of each subject. The visual cortex was morphologically variable among the subjects, but the P100m dipoles were all localized at the lateral bottom of the calcarine fissure. Moreover, these P100m dipoles had similar orientations for both half- or quadrant-field stimuli. These results suggest that the P100m is located in a smaller part of the striate cortex than previously reported.     

Trajectories of shifting of dipole sources of visual evoked potentials over the human brain

Dynamic study of 3D localization of the equivalent current dipole (ECD) sources of visual evoked potentials (EP) in the human brain was performed in 18 healthy subjects using a two-dipole model. Dipole tracing was performed for relatively early EP components (N1, P1, and N2) with 1-ms step. The analysis confirmed localization of these ECDs mainly in the right occipital cortex and revealed their successive shift over this area in the anterior-medial direction and then backwards in all subjects during generation of the EP components. Typically, some successive arch-like trajectories of the shift were revealed (75.8%); their duration was relatively standard (about 25 ms) and did not depend on the stimulus shape and EP phase. Between the 1st and the 2nd trajectories (110-120 ms after the stimulus onset) a jump in ECD coordinates in the medial direction was found in 85% of cases. Possible significance of the findings for the insight into dynamic topography of the visual feature processing in the human brain is discussed.     

Delayed striate cortical activation during spatial attention

Recordings of event-related potentials (ERPs) and event-related magnetic fields (ERMFs) were combined with functional magnetic resonance imaging (fMRI) to study visual cortical activity in humans during spatial attention. While subjects attended selectively to stimulus arrays in one visual field, fMRI revealed stimulus-related activations in the contralateral primary visual cortex and in multiple extrastriate areas. ERP and ERMF recordings showed that attention did not affect the initial evoked response at 60-90 ms poststimulus that was localized to primary cortex, but a similarly localized late response at 140-250 ms was enhanced to attended stimuli. These findings provide evidence that the primary visual cortex participates in the selective processing of attended stimuli by means of delayed feedback from higher visual-cortical areas.     

Dipole models of alpha-rhythm generators

Different hypothesis of the alpha rhythm origin were tested by dipole simulation of the alpha rhythm sources. EEG was recorded during driving photic stimulation in order to increase the signal-to-noise ratio. Models with fixed and moving dipoles were analyzed. Dipole sources were compared with magnetic resonance imaging (MRI) scans to find the exact location of oscillations in brain anatomical structures. A two-level multiple dipole model was found to fit the EEG most adequately. The first level was represented by two oscillators localized in the thalamic reticular nuclei, and the second level is associated with two modality-specific oscillators localized in the respective cortical areas.     

Brain potentials associated with pattern displacement and saccadic eye movements in humans and rhesus monkeys

Visual evoked potentials (VEPs) to the onset of motion of visual patterns and brain responses associated with saccadic eye movements (SRPs) were compared in human subjects and in rhesus monkeys. Three different velocities of pattern motion were employed. In humans, brain responses were recorded from six scalp areas. In monkeys, transcortical recordings were obtained from chronically implanted electrodes in the occipital, temporo-parietal, and frontal areas. In humans there was a clear difference in VEPs to the pattern motion between the anterior (Fz, Cz) and posterior (Pz, Oz) scalp regions. The earliest component was a positive peak at 85 ms at Oz followed by a negativity around 110 ms. In the fronto-central leads the VEP was characterized by a negativity at 145 ms and a subsequent broad positive component around 250 ms. SRP responses differed in the early components from the VEPs to pattern motion but a good correspondence was found in the morphology of the late components of the two types of brain potentials. Furthermore, flashed-on VEPs and SRPs elicited a late positivity of more pronounced amplitude than VEPs to pattern displacement. In monkeys similar findings were found: an early negative component of the pattern-displacement VEP could not be observed in the SRP responses over the visual cortex while the late portion of the SRP waveform was greater than the late positivity of the VEP to motion-onset.     

Temporal and topographic characteristics of evoked potentials in the conflict of two consecutive visual stimuli in a working memory task

Behavioral reactions and brain mechanisms involved in processing two matching or mismatching (conflicting) visual stimuli were studied in healthy subjects (mean age 22.57 ± 0.46 years). Line orientations (vertical, horizontal, or 45°) were used as stimuli and were presented with an interval of 1500–1800 ms. The reaction time was shown to increase in the case of a conflict of two orientations as compared with matching orientations. The reaction time depended on the orientation of the reference stimulus and was minimal when a vertical line was used as a reference. An increase in N2 negativity (time window 200–280 ms) in the frontal and parietal cortical areas was identified as an informative indicator of a conflict between the current orientation and the orientation stored in working memory. The dipole sources of N2 were localized to the prefrontal cortex (middle frontal gyrus, frontal pole, and pars orbitalis). The N2 amplitude was found to depend on the orientation of the first stimulus in a pair, being higher in the case of a 45° orientation. The visual areas were shown to play a role in detecting a conflict of two consecutive signals because the early sensory components increased in amplitude. The results implicate cortical structures, including the sensory-specific visual, parietal, and prefrontal areas, in comparing consecutive visual signals and detecting their conflict.     

Source modeling of esophageal evoked potentials

Evoked potentials can be recorded from the scalp after stimulation of the esophagus by balloon distension. The purpose of this study was to estimate the number and localization of sources contributing to the esophageal evoked potential (EEP). The EEP was recorded from 32 scalp electrodes in 5 healthy subjects. Spatio-temporal dipole modeling was performed in the time interval from 185 msec to 525 msec after stimulation (mean values). The EEP was best explained by the combined activity of 1 dipole located relatively high in the midline and 2 lateral dipoles. Given the anatomical projection of esophageal sensory fibers and the location of these dipoles, the sources were probably located in the cingulate gyri and insular cortex. There was no evidence that sources in the lower brain-stem contributed to the scalp recorded EEP.     

Rhythmic photostimulation and a number of alpha-rhythm dipoles in the human brain

Dynamic study of the equivalent current dipoles (ECDs) of alpha rhythm in the human brain was performed in a one-dipole model. The number of ECDs was shown to be dependent on the time course of visual stimulation, on the phase shift of triggering of flickers with alpha-rhythm frequency from alpha-wave, as well as on the type of visual illusions produced in subjects by this stimulation. The data are discussed in accordance with the "scanning hypothesis" that predict a certain functional meaning of the spreading alpha-wave for cortical processing of sensory information in the human brain.     

形状识别的功能定位和时间过程:功能磁共振与脑电结合的研究

通过结合具有高空间分辨率的功能磁共振成像（fMRI）和具有高时间分辨率的128导脑电事件相关电位（ERP）两项技术,测量了视皮层腹侧区域对图形形状识别任务反应的空间定位和时间过程。fMRI的实验结果表明,图形的形状和觉引起了腹测GTi／GF皮层区域的兴奋。进一步,基于fMRI兴奋区域的种子偶极子模型拟合的的ERP动态定位分析的结果和自由运动的偶极子模型拟合的ERP定位分析结果表明：GTi／GF区域活动的时间发生在刺激呈现之后132－176ms时间段,峰值150ms左右,相应于ERP的N1成分。这些结果在人类大脑皮层上同时确定了视觉通路中涉及图形形状识别的兴奋区域和兴奋的时间过程。     

Dynamic construction of stimulus values in the ventromedial prefrontal cortex

Signals representing the value assigned to stimuli at the time of choice have been repeatedly observed in ventromedial prefrontal cortex (vmPFC). Yet it remains unknown how these value representations are computed from sensory and memory representations in more posterior brain regions. We used electroencephalography (EEG) while subjects evaluated appetitive and aversive food items to study how event-related responses modulated by stimulus value evolve over time. We found that value-related activity shifted from posterior to anterior, and from parietal to central to frontal sensors, across three major time windows after stimulus onset: 150-250 ms, 400-550 ms, and 700-800 ms. Exploratory localization of the EEG signal revealed a shifting network of activity moving from sensory and memory structures to areas associated with value coding, with stimulus value activity localized to vmPFC only from 400 ms onwards. Consistent with these results, functional connectivity analyses also showed a causal flow of information from temporal cortex to vmPFC. Thus, although value signals are present as early as 150 ms after stimulus onset, the value signals in vmPFC appear relatively late in the choice process, and seem to reflect the integration of incoming information from sensory and memory related regions.     

Localization of human brain areas activated for chaotic and ordered pattern perception

The aim of our work was to localize cortical areas involved in the processing of incomplete figures using functional MRI (fMRI) for 8 healthy volunteers (18-30 year old) with the did of anatomical and fMRI fast imaging technique: echo planar imaging (EPI), whole brain scan (36 slices) matrix 64 x 64, 3.7 second. We used 1.5 T MR-scanner and BOLD-method (Blood Oxygenation Level Dependent), based on distinctions of magnetic properties of hemoglobin. Fast imaging technique on modern MR-scanners with > or = 1.5 T provides precise statistical maps of oxygenation increase with high spatial resolution. For test stimuli we used matrix of Gabor grating. We used two types of 10 x 10 matrices with chaotic and ordered orientation of Gabor gratings. The size, brightness and contrast of the stimuli were identical. The chaotic and ordered patterns activated different brain areas. We establish that ordered patterns activated only primary visual cortex - V1 and V2, (BA17-18), wheareas chaotic patterns activated in addition primary visual cortex, the V3,V4,V5 (BA19) of the occipital cortex and the area 7 of parietal area (BA7) classification. Decision making for that task is localized in prefrontal and frontal cortex, including (BA 6, 9, 10).     

Role of filtration in localization of the evoked potential dipoles in the human brain: experiment and simulation

3D dipole tracing with 1 ms step of visual evoked potentials recorded from 40 electrodes was performed under exposition of crosses in 5 healthy human subjects. The data on dipole displacement were compared with prediction of the simulation study on distortion of dipole localization by the signal filtration in the low-frequency band. These predictions were experimentally confirmed: the effect depends on the degree of filtration (0.1 or 0.5 Hz) and on the latency of EP waves. Localization and strength of P1 dipoles were not changed under filtration, while for later components--N2 and especially P3--they changed significantly. For the improvement of these distortions time constant of the amplification tract must be some times longer than the time of the dipole activity.     

Effects of cavities on EEG dipole localization and their relations with surface electrode positions

Effects of cavities in the human head on EEG dipole localization have been investigated by computer simulation. The human head is represented by a homogeneous spherical conductor including an eccentric spherical cavity which approximates effects of actual cavities inside the head. The homogeneous sphere model is used for assessing the effects caused by neglecting the cavity in the volume conductor model in the inverse dipole fitting procedure. Four electrode configurations have been examined to investigate their relation to the EEG inverse dipole solution. After examination of 2520 dipoles in the brain, the effects of cavities in the human head are found to be negligible when the dipole is located in the cortex or in the subcortex. When the dipole is located in the brain stem, the EEG inverse dipole solution is strongly affected by the cavity and is sensitive to the electrode configuration on the scalp. The EEG inverse dipole solution in the deep brain is sensitive to inhomogeneity in the lower part of the head when a single positive or negative potential pole is observed by the electrodes on the scalp, and at the same time is sensitive to the extent of the scalp covered by the electrodes. In conclusion, the electrodes should cover as much of the upper scalp as possible for deep source localization.     

Neural Correlates of Central Inhibition during Physical Fatigue

Central inhibition plays a pivotal role in determining physical performance during physical fatigue. Classical conditioning of central inhibition is believed to be associated with the pathophysiology of chronic fatigue. We tried to determine whether classical conditioning of central inhibition can really occur and to clarify the neural mechanisms of central inhibition related to classical conditioning during physical fatigue using magnetoencephalography (MEG). Eight right-handed volunteers participated in this study. We used metronome sounds as conditioned stimuli and maximum handgrip trials as unconditioned stimuli to cause central inhibition. Participants underwent MEG recording during imagery of maximum grips of the right hand guided by metronome sounds for 10 min. Thereafter, fatigue-inducing maximum handgrip trials were performed for 10 min; the metronome sounds were started 5 min after the beginning of the handgrip trials. The next day, neural activities during imagery of maximum grips of the right hand guided by metronome sounds were measured for 10 min. Levels of fatigue sensation and sympathetic nerve activity on the second day were significantly higher relative to those of the first day. Equivalent current dipoles (ECDs) in the posterior cingulated cortex (PCC), with latencies of approximately 460 ms, were observed in all the participants on the second day, although ECDs were not identified in any of the participants on the first day. We demonstrated that classical conditioning of central inhibition can occur and that the PCC is involved in the neural substrates of central inhibition related to classical conditioning during physical fatigue.     

Perceptual echoes at 10 Hz in the human brain

The occipital alpha rhythm (～10 Hz) is the most prominent electrophysiological activity in the awake human brain, yet its functional role and relation to visual perception are little understood. Transient stimuli normally elicit a short series of positive and negative deflections lasting between 300 and 500 ms: the visual-evoked potential (VEP). Alpha oscillations, on the other hand, are generally suppressed by transient visual input; they only augment in response to periodic ("steady-state") inputs around 10 Hz. Here, we applied reverse-correlation techniques to the visual presentation of random, nonperiodic dynamic stimulation sequences and found that the brain response to each stimulus transient was not merely a short-lived VEP but also included a strong ～10 Hz oscillation that lasted for more than 1 s. In other words, the alpha rhythm implements an "echo" or reverberation of the input sequence. These echoes are correlated in magnitude and frequency with the observer's occipital alpha rhythm, are enhanced by visual attention, and can be rendered perceptually apparent in the form of ～10 Hz flicker. These findings suggest a role for the alpha rhythm in the maintenance of sensory representations over time.     

Computation of color and brightness differences by neurons in the rabbit visual cortex

Changes in activity of 54 neurons in the rabbit visual cortex evoked by the replacement of eight color and eight achromatic stimuli in pairs were analyzed. The diffused stimuli generated by color SVGA monitor were used in the experiments. The earliest response of phasic neurons (50-90 ms after the replacement) was strongly correlated with differences between stimuli in color or intensity. This response ("the signal of differences") was used as a basis of a matrix (8 x 8) constructed for each neuron. Such matrices included mean numbers of spikes per second in responses to changes of different stimuli pairs. All matrices were subjected to factor analysis, and the basic axes (the main factors) of sensory spaces were revealed. It was found that 16 neurons (30%) detected only achromatic differences between stimuli. Perceptual spaces of these neurons were two-dimensional with brightness and darkness orthogonal axes. The spaces of 12 neurons (22%) were four-dimensional with two chromatic and two achromatic axes. The structure of the perceptual space reconstructed from neuronal spikes was similar to the space calculated from the early VEP components recorded under similar conditions and to another space reconstructed on the basis of rabbit's instrumental learning. The fundamental coincidence of color spaces revealed by different methods may reflect the general principle of vector coding in the visual system and suggests the coexistence of two independent cortical mechanisms of the detection of chromatic and achromatic differences.     

The influence of conductivity changes in boundary element compartments on the forward and inverse problem in electroencephalography and magnetoencephalography.

Source localization based on magnetoencephalographic and electroencephalographic data requires knowledge of the conductivity values of the head. The aim of this paper is to examine the influence of compartment conductivity changes on the neuromagnetic field and the electric scalp potential for the widely used three compartment boundary element models. Both the analysis of measurement data and the simulations with dipoles distributed in the brain produced two significant results. First, we found the electric potentials to be approximately one order of magnitude more sensitive to conductivity changes than the magnetic fields. This was valid for the field and potential topology (and hence dipole localization), and for the amplitude (and hence dipole strength). Second, changes in brain compartment conductivity yield the lowest change in the electric potentials topology (and hence dipole localization), but a very strong change in the amplitude (and hence in the dipole strength). We conclude that for the magnetic fields the influence of compartment conductivity changes is not important in terms of dipole localization and strength estimation. For the electric potentials however, both dipole localization and strength estimation are significantly influenced by the compartment conductivity.     

Attention modulates earliest responses in the primary auditory and visual cortices

A fundamental question about the neural correlates of attention concerns the earliest sensory processing stage that it can affect. We addressed this issue by recording magnetoencephalography (MEG) signals while subjects performed detection tasks, which required employment of spatial or nonspatial attention, in auditory or visual modality. Using distributed source analysis of MEG signals, we found that, contrary to previous studies that used equivalent current dipole (ECD) analysis, spatial attention enhanced the initial feedforward response in the primary visual cortex (V1) at 55-90 ms. We also found attentional modulation of the putative primary auditory cortex (A1) activity at 30-50 ms. Furthermore, we reproduced our findings using ECD modeling guided by the results of distributed source analysis and suggest a reason why earlier studies using ECD analysis failed to identify the modulation of earliest V1 activity.     

Neural basis of the ventriloquist illusion

The ventriloquist creates the illusion that his or her voice emerges from the visibly moving mouth of the puppet [1]. This well-known illusion exemplifies a basic principle of how auditory and visual information is integrated in the brain to form a unified multimodal percept. When auditory and visual stimuli occur simultaneously at different locations, the more spatially precise visual information dominates the perceived location of the multimodal event. Previous studies have examined neural interactions between spatially disparate auditory and visual stimuli [2-5], but none has found evidence for a visual influence on the auditory cortex that could be directly linked to the illusion of a shifted auditory percept. Here we utilized event-related brain potentials combined with event-related functional magnetic resonance imaging to demonstrate on a trial-by-trial basis that a precisely timed biasing of the left-right balance of auditory cortex activity by the discrepant visual input underlies the ventriloquist illusion. This cortical biasing may reflect a fundamental mechanism for integrating the auditory and visual components of environmental events, which ensures that the sounds are adaptively localized to the more reliable position provided by the visual input.