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.     

Dipole tracing of visual evoked potentials in human brain

3D tracing of equivalent current dipoles (ECDs) of averaged human visual evoked potentials (VEP) by their distribution across a 34-electrode array was obtained under short presentation of pattern-onset stimuli (sets of 45 horizontal, vertical bars or crosses). Using a 2-dipole spherical three-layer model, we dynamically (step of 1 ms) localized dipoles in four healthy subjects. Dipole locations were matched to anatomical brain regions visualized in structural MRI. Best-fitting source parameters were superimposed on MR images of each subject to identify the anatomical structures giving rise to the surface patterns. It was found that during 50-300 ms following the onset of the stimuli, the ECDs in all subjects were localized in the occipital cortex and demonstrated reliable systematic shift in localization. Two local (1-2 cm3) zones of the preferable dipole attendance were found at 5-6 cm behind zero line: the first one was localized near the midline of the brain, whereas the other zone was situated in the right hemisphere at a distance of 6-7 cm from the first zone. Their localization and strength of activation were reliably different for crosses and lines and changed during VEP generation. Zones of relatively rare dipole attendance were found also. The data are discussed in relation to localization of initial and endpoint of ECDs trajectories, as well as with sensitivity of the visual cortex to line crossing and branching.     

Functionally independent components of early event-related potentials in a visual spatial attention task.

Spatial visual attention modulates the first negative-going deflection in the human averaged event-related potential (ERP) in response to visual target and non-target stimuli (the N1 complex). Here we demonstrate a decomposition of N1 into functionally independent subcomponents with functionally distinct relations to task and stimulus conditions. ERPs were collected from 20 subjects in response to visual target and non-target stimuli presented at five attended and non-attended screen locations. Independent component analysis, a new method for blind source separation, was trained simultaneously on 500 ms grand average responses from all 25 stimulus-attention conditions and decomposed the non-target N1 complexes into five spatially fixed, temporally independent and physiologically plausible components. Activity of an early, laterally symmetrical component pair (N1aR and N1aL) was evoked by the left and right visual field stimuli, respectively. Component N1aR peaked ca. 9 ms earlier than N1aL. Central stimuli evoked both components with the same peak latency difference, producing a bilateral scalp distribution. The amplitudes of these components were no reliably augmented by spatial attention. Stimuli in the right visual field evoked activity in a spatio-temporally overlapping bilateral component (N1b) that peaked at ca. 180 ms and was strongly enhanced by attention. Stimuli presented at unattended locations evoked a fourth component (P2a) peaking near 240 ms. A fifth component (P3f) was evoked only by targets presented in either visual field. The distinct response patterns of these components across the array of stimulus and attention conditions suggest that they reflect activity in functionally independent brain systems involved in processing attended and unattended visuospatial events.     

Ontogenetic characteristics of the development of slow negative and positive potentials during the performance of a visual perceptive task

In age aspect, topography and parameters were studied of late EPs and slow negative components (CNV) to successively presented visual stimuli: the preliminary signal and two compared contour images. Age characteristics of development of the frontal negativity N450 and the late positive complex (LPC) were revealed. in children 7 and 10 years old, the component N450 was present in response both to the preliminary and the first stimulus in the compared pair; LPC was recorded to all three presented visual stimuli. In 17 years old subjects and in adults, selectivity of N450 formation in the EP to the first of the compared stimuli and LPC to the second (imperative) stimulus correlates with involvement of rostral brain areas--central in youths, central and frontal ones in adults--both in the process of waiting (CNV) and evaluation of coming information (LPC).     

Amplitude-temporal parameters of the evoked potentials of the visual and motor areas in the visual perception and mental representation of an image

Amplitude-temporal analysis was carried out of the EP components of the visual and motor areas elicited by neutral (diffuse light) and structural (checker board pattern) stimuli in different situations, defined by instruction. Interserial comparisons showed that at any instruction, the latency of the first EP component of the motor areas is reduced; as a result it can appear here simultaneously with the EP of the visual areas. At the instruction involving the subject in the process of active change of perception, activation of the right hemisphere, including the motor area, is manifest by EP parameters, while the right occipital area is activated in response to the structural stimulus, and the left one--in response to the neutral stimulus. At complication of the stimulus or instruction, the period is prolonged when the latency of EP components of the motor area is shorter than the latency of the isopolar components of the visual area--from 120 to 150 ms in response to the neutral stimuli and the neutral with their counting; from 90 to 150 ms in response to the structural stimuli; from 80 to 210 ms in response to the neutral stimuli with mental representation of the structural one.     

Functional aspects of evoked alpha and theta responses in humans and cats

A systems theoretical approach was used to compare possible functional roles of theta (4--7 Hz) and alpha (8--15 Hz) response components of brain evoked potentials. These response components were described earlier by Ba\c sar (1980). We recorded EEG and evoked potentials (EPs) from occipital scalp locations in 11 subjects. We used auditory and visual stimuli as inadequate and adequate stimuli, respectively (``cross-modality' measurements). The combined EEG-EP epochs were analysed in frequency domain with fast Fourier transform and adaptive digital filters. Alpha (8--15 Hz) response components turned out to be dependent on whether the stimulus was adequate or not (median amplitude with inadequate vs. adequate stimulation: vs. ). Theta (4--7 Hz) response components were less dependent on stimulus modality (inadequate vs. adequate stimulation: vs. ). In EP recordings the occipital alpha response almost disappeared in the first 250 ms following auditory stimulation. Comparable behaviour was observed in similar experiments with recordings from the cat visual cortex (area 17) and with occipital magnetoencephalographic recordings. Taking into account the above-mentioned previous reports on intracranial recordings in primary sensory areas of the cat brain and preliminary results of magnetoencephalographic measurements, we propose the following hypothesis: alpha responses in a time window of about 250 ms after stimulation might predominantly reflect primary sensory processing whereas the theta responses in the first 250 ms after stimulation might be more involved in supra-modality -- or cross-modality -- associative-cognitive processing. Received: 25 February 1994 / Accepted in revised form: 5 August 1994     

Visual Mislocalization of Moving Objects in an Audiovisual Event

The present study investigated the influence of an auditory tone on the localization of visual objects in the stream/bounce display (SBD). In this display, two identical visual objects move toward each other, overlap, and then return to their original positions. These objects can be perceived as either streaming through or bouncing off each other. In this study, the closest distance between object centers on opposing trajectories and tone presentation timing (none, 0 ms, ± 90 ms, and ± 390 ms relative to the instant for the closest distance) were manipulated. Observers were asked to judge whether the two objects overlapped with each other and whether the objects appeared to stream through, bounce off each other, or reverse their direction of motion. A tone presented at or around the instant of the objects’ closest distance biased judgments toward “non-overlapping,” and observers overestimated the physical distance between objects. A similar bias toward direction change judgments (bounce and reverse, not stream judgments) was also observed, which was always stronger than the non-overlapping bias. Thus, these two types of judgments were not always identical. Moreover, another experiment showed that it was unlikely that this observed mislocalization could be explained by other previously known mislocalization phenomena (i.e., representational momentum, the Fröhlich effect, and a turn-point shift). These findings indicate a new example of crossmodal mislocalization, which can be obtained without temporal offsets between audiovisual stimuli. The mislocalization effect is also specific to a more complex stimulus configuration of objects on opposing trajectories, with a tone that is presented simultaneously. The present study promotes an understanding of relatively complex audiovisual interactions beyond simple one-to-one audiovisual stimuli used in previous studies.     

基于小波变换的单次诱发电位信号时频分析

比较小波变换和平均叠加两种方法提取“模拟自然阅读”刺激模式下的诱发脑电信号,分析其时频特性,并进行脑功能源分布定位分析。结果显示,采用平均叠加法来提取和分析诱发电位信号,损失了某些重要的诱发电位成分,且其功能源分布定位反映的只是等效功能源的静态过程;而使用小波变换和脑功能源定位来提取和分析单次诱发电位信号,既能观察到丰富的诱发电位成分,又能反映脑功能源的实时动态活动过程。这表明,小波变换下的时频分析是脑电信号处理的一种可行的新方法。     

Pre-stimulus spectral EEG patterns and the visual evoked response

The relationship between the latencies and amplitudes of the N1 and P2 components of the visual evoked potential (VEP) and the psychophysiological state of the brain immediately preceding the time of the stimulus has been investigated in 7 male subjects. Power spectral measures in the delta, theta, alpha and beta bands of the 1 sec pre-stimulus EEG were used to assess the brain state, and low intensity flashes, delivered randomly between 2 and 6 whole seconds, were used as the stimuli. Trials were ranked separately according to the relative amounts of pre-stimulus power in each EEG band and were partitioned into groups by an equal pre-stimulus spectral power criterion. Averaged EPs were computed from these groups and multiple regression analysis was used to relate pre-stimulus spectral power values to EP features. Five of the 7 subjects displayed consistent increases in N1-P2 amplitude as a function of increasing pre-stimulus relative alpha power. The between-subjects effect of pre-stimulus EEG on N1 latency was small, but was moderate for P2 latency (both significant). Both N1 and P2 latency were found to decrease with increasing amounts of pre-stimulus relative delta and theta power.     

视觉空间注意事件相关电位的协同学分析

在用事件相关电位(event-related potentials,ERP)研究视觉空间注意问题时,直接观察ERP数据就可得出,空间注意的主要作用是对视觉信息处理的调制,它出现在刺激开始后大约80～250ms,主要表现为枕叶的P1、N1和P2波有明显的增强但它们的潜伏期没有变化。采用基于协同学的时空模式分解方法,把视觉空间注意ERP分解为3个模式成分。结果表明,注意不仅使模式1的第一个正波成分(P11)、第一个负波成分(N11)以及第二个正波成分(P12)增强,还使模式3的第一个正波成分(P31)的潜伏期缩短。用探照灯模型对这些现象作了初步解释,说明该方法是研究注意ERP的一种有潜力的新方法。     

Evoked potentials of the cerebral cortex during the comparison of visual stimuli

Visual evoked potentials (EP) were recorded when the test subjects accomplished the tasks of a comparison of a current stimulus with the previous one, the stimuli being presented in a continuous sequence. In the first task, rare repetition of two stimuli (Russian letters) in the continuously changing flow of stimuli was relevant, and the test subject had to press the button when it happened; in the second task, the relevant stimulus was a rare change in the flow of stimuli. The influence of the stimulus repetition/change factor on EP was analyzed. The processes related to the comparison of the current and previous stimuli were most manifest in four time intervals: 120–140, 180–210, 260–280, and 350–370 ms. The occipito-temporal component of EP revealed in the interval of 180–210 ms, which we denoted as the negative component of visual mismatch (NCVM), proved a special component, differing in its functional and temporal characteristics from theN _2b component. WhereasN _2b is modulated by the factor of stimulus probability, the NCVM by that of stimulus repetition/change.     

Motion-reversal visual evoked responses.

Motion-reversal visual evoked responses (VERs) have remarkable waveform variability. In our opinion this is caused by the alternative predominance of either motion or pattern-onset/offset related components. The motion dependent component of motion-reversal VER closely resembles motion-onset VER (main negative peak with the latency of about 170 ms), the first positive peak (with the latency of about 100 ms) corresponds to the pattern-onset component and the second non-constant positive peak (with the latency of about 130 ms) seems to be identical with the pattern-offset positivity. The differences in expression of these components are dependent on some stimulus characteristics (mainly on the contrast of a structure, velocity of motion, retinal localization of the stimulus) and on substantial differences in the sensitivity of subjects to motion stimulation.     

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.     

Gustatory evoked cortical activity in humans studied by simultaneous EEG and MEG recording

Evoked potentials are widely used in clinical medicine for objective evaluation of sensory disturbances. However, gustatory evoked potentials (GEPs) have not been extensively studied due to lack of agreement among investigators regarding the waveforms. In this study GEPs and gustatory magnetic fields (GEMfs) were simultaneously recorded from five subjects in response to 0.3 M NaCl in an attempt to establish GEP recording as an objective gustatometer. Each subject received a total of 240 stimulus presentations over six sessions. Three GEP components (P1, N1 and P2) were observed and correlated with their corresponding equivalent current dipoles (ECD1, ECD2 and ECD3, respectively). ECD1 was localized to area G in all subjects, P1 being the indicator of intact gustatory projection to area G. No significant GEP activity was detected during the time preceding P1, which suggests that there was no activity in cortical gyri other than that detected by magnetoencephalography. ECD2 and ECD3 were localized to various cortical structures, including the inferior insula and the superior temporal sulcus, indicating that N1 and P2 reflect higher order gustatory functions. The present results indicate that measurement of GEPs may be useful for objective evaluation of gustatory disturbance.     

Evidence for dissociation between the perceptual and visuomotor systems in humans

When a visual stimulus is continuously moved behind a small stationary window, the window appears displaced in the direction of motion of the stimulus. In this study we showed that the magnitude of this illusion is dependent on (i) whether a perceptual or visuomotor task is used for judging the location of the window (ii) the directional signature of the stimulus, and (iii) whether or not there is a significant delay between the end of the visual presentation and the initiation of the localization measure. Our stimulus was a drifting sinusoidal grating windowed in space by a stationary, two-dimensional, Gaussian envelope (sigma=1 cycle of sinusoid). Localization measures were made following either a short (200 ms) or long (4.2 s) post-stimulus delay. The visuomotor localization error was up to three times greater than the perceptual error for a short delay. However, the visuomotor and perceptual localization measures were similar for a long delay. Our results provide evidence in support of the hypothesis that separate cortical pathways exist for visual perception and visually guided action and that delayed actions rely on stored perceptual information.     

Human orienting reaction: the role of genetic and environmental factors in the variability of evoked potentials and autonomic components

The effect of genetic and environmental factors on interindividual differences in GSR, heart rate and EP components of orienting reaction to neutral acoustical stimulus and to the same stimulus at a very high intensity was analysed in 22 pairs of monozygotic and 21 pairs of dizygotic twins. Genetic factors appeared to be important for the variability of all GSR characteristics, N100 and P180 EP amplitudes, magnitude of the HR response to the strong stimulus and HR habituation to the weak stimulus. Environmental factors influenced greatly the variances of late EP components but only in response to the weak stimulus. Different role of genetic and environmental factors in motivation and cognitive components of orienting reaction points to the involvement of several relatively independent functional systems in it.     

Stimulus timing-dependent plasticity in cortical processing of orientation.

The relative timing of presynaptic and postsynaptic spikes plays a critical role in activity-induced synaptic modification. Here we examined whether plasticity of orientation selectivity in the visual cortex depends on stimulus timing. Repetitive pairing of visual stimuli at two orientations induced a shift in orientation tuning of cat cortical neurons, with the direction of the shift depending on the temporal order of the pair. Induction of a significant shift required that the interval between the pair fall within +/-40 ms, reminiscent of the temporal window for spike timing-dependent synaptic plasticity. Mirroring the plasticity found in cat visual cortex, similar conditioning also induced a shift in perceived orientation by human subjects, further suggesting functional relevance of this phenomenon. Thus, relative timing of visual stimuli can play a critical role in dynamic modulation of adult cortical function, perhaps through spike timing-dependent synaptic plasticity.     

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.     

The extrastriate generators of the EP to checkerboard onset. A source localization approach

The cortical origin of the pattern onset EP has been investigated over a time window which covers the entire positive-negative-positive complex of the pattern onset EP. On the basis of a dipole source localization approach, the position, orientation and strength of the underlying sources of the pattern onset EP were estimated. For large check stimuli, chosen to have a weak edge specific component in the response, still two components are needed to account for the variance of the responses. Each component corresponds to a single dipole source, and both originate in the extrastriate cortex. These components dominate, respectively, the initial and the late positive peaks of the pattern onset EP.The equivalent dipole sources of the two components show different behaviors with respect to the position of the stimulus in the visual field. The topography and behavior of the equivalent dipole source underlying the early positive component suggest an origin in area 18. The invariance with stimulus location of the dipole source underlying the late positive component suggests an origin beyond area 18. The different topographies of the components also account for the differences in surface distribution of the pattern onset EP to large check stimulation of the upper and lower sectors of the visual field.     

Influence of Visual Motion,Suggestion, and Illusory Motion on Self-Motion Perception in the Horizontal Plane

A moving visual field can induce the feeling of self-motion or vection. Illusory motion from static repeated asymmetric patterns creates a compelling visual motion stimulus, but it is unclear if such illusory motion can induce a feeling of self-motion or alter self-motion perception. In these experiments, human subjects reported the perceived direction of self-motion for sway translation and yaw rotation at the end of a period of viewing set visual stimuli coordinated with varying inertial stimuli. This tested the hypothesis that illusory visual motion would influence self-motion perception in the horizontal plane. Trials were arranged into 5 blocks based on stimulus type: moving star field with yaw rotation, moving star field with sway translation, illusory motion with yaw, illusory motion with sway, and static arrows with sway. Static arrows were used to evaluate the effect of cognitive suggestion on self-motion perception. Each trial had a control condition; the illusory motion controls were altered versions of the experimental image, which removed the illusory motion effect. For the moving visual stimulus, controls were carried out in a dark room. With the arrow visual stimulus, controls were a gray screen. In blocks containing a visual stimulus there was an 8s viewing interval with the inertial stimulus occurring over the final 1s. This allowed measurement of the visual illusion perception using objective methods. When no visual stimulus was present, only the 1s motion stimulus was presented. Eight women and five men (mean age 37) participated. To assess for a shift in self-motion perception, the effect of each visual stimulus on the self-motion stimulus (cm/s) at which subjects were equally likely to report motion in either direction was measured. Significant effects were seen for moving star fields for both translation (p = 0.001) and rotation (p<0.001), and arrows (p = 0.02). For the visual motion stimuli, inertial motion perception was shifted in the direction consistent with the visual stimulus. Arrows had a small effect on self-motion perception driven by a minority of subjects. There was no significant effect of illusory motion on self-motion perception for either translation or rotation (p>0.1 for both). Thus, although a true moving visual field can induce self-motion, results of this study show that illusory motion does not.