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S Makeig

Publications and source records attributed to S Makeig.

At least 19 recordsLinked to original sources

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.

Adolescent

Functionally independent components of the late positive event-related potential during visual spatial attention.

Human event-related potentials (ERPs) were recorded from 10 subjects presented with visual target and nontarget stimuli at five screen locations and responding to targets presented at one of the locations. The late positive response complexes of 25-75 ERP average waveforms from the two task conditions were simultaneously analyzed with Independent Component Analysis, a new computational method for blindly separating linearly mixed signals. Three spatially fixed, temporally independent, behaviorally relevant, and physiologically plausible components were identified without reference to peaks in single-channel waveforms. A novel frontoparietal component (P3f) began at approximately 140 msec and peaked, in faster responders, at the onset of the motor command. The scalp distribution of P3f appeared consistent with brain regions activated during spatial orienting in functional imaging experiments. A longer-latency large component (P3b), positive over parietal cortex, was followed by a postmotor potential (Pmp) component that peaked 200 msec after the button press and reversed polarity near the central sulcus. A fourth component associated with a left frontocentral nontarget positivity (Pnt) was evoked primarily by target-like distractors presented in the attended location. When no distractors were presented, responses of five faster-responding subjects contained largest P3f and smallest Pmp components; when distractors were included, a Pmp component appeared only in responses of the five slower-responding subjects. Direct relationships between component amplitudes, latencies, and behavioral responses, plus similarities between component scalp distributions and regional activations reported in functional brain imaging experiments suggest that P3f, Pmp, and Pnt measure the time course and strength of functionally distinct brain processes.

Adult

Spatially independent activity patterns in functional MRI data during the stroop color-naming task.

A method is given for determining the time course and spatial extent of consistently and transiently task-related activations from other physiological and artifactual components that contribute to functional MRI (fMRI) recordings. Independent component analysis (ICA) was used to analyze two fMRI data sets from a subject performing 6-min trials composed of alternating 40-sec Stroop color-naming and control task blocks. Each component consisted of a fixed three-dimensional spatial distribution of brain voxel values (a "map") and an associated time course of activation. For each trial, the algorithm detected, without a priori knowledge of their spatial or temporal structure, one consistently task-related component activated during each Stroop task block, plus several transiently task-related components activated at the onset of one or two of the Stroop task blocks only. Activation patterns occurring during only part of the fMRI trial are not observed with other techniques, because their time courses cannot easily be known in advance. Other ICA components were related to physiological pulsations, head movements, or machine noise. By using higher-order statistics to specify stricter criteria for spatial independence between component maps, ICA produced improved estimates of the temporal and spatial extent of task-related activation in our data compared with principal component analysis (PCA). ICA appears to be a promising tool for exploratory analysis of fMRI data, particularly when the time courses of activation are not known in advance.

Algorithms

Analysis of fMRI data by blind separation into independent spatial components.

Current analytical techniques applied to functional magnetic resonance imaging (fMRI) data require a priori knowledge or specific assumptions about the time courses of processes contributing to the measured signals. Here we describe a new method for analyzing fMRI data based on the independent component analysis (ICA) algorithm of Bell and Sejnowski ([1995]: Neural Comput 7:1129-1159). We decomposed eight fMRI data sets from 4 normal subjects performing Stroop color-naming, the Brown and Peterson work/number task, and control tasks into spatially independent components. Each component consisted of voxel values at fixed three-dimensional locations (a component "map"), and a unique associated time course of activation. Given data from 144 time points collected during a 6-min trial, ICA extracted an equal number of spatially independent components. In all eight trials, ICA derived one and only one component with a time course closely matching the time course of 40-sec alternations between experimental and control tasks. The regions of maximum activity in these consistently task-related components generally overlapped active regions detected by standard correlational analysis, but included frontal regions not detected by correlation. Time courses of other ICA components were transiently task-related, quasiperiodic, or slowly varying. By utilizing higher-order statistics to enforce successively stricter criteria for spatial independence between component maps, both the ICA algorithm and a related fourth-order decomposition technique (Comon [1994]: Signal Processing 36:11-20) were superior to principal component analysis (PCA) in determining the spatial and temporal extent of task-related activation. For each subject, the time courses and active regions of the task-related ICA components were consistent across trials and were robust to the addition of simulated noise. Simulated movement artifact and simulated task-related activations added to actual fMRI data were clearly separated by the algorithm. ICA can be used to distinguish between nontask-related signal components, movements, and other artifacts, as well as consistently or transiently task-related fMRI activations, based on only weak assumptions about their spatial distributions and without a priori assumptions about their time courses. ICA appears to be a highly promising method for the analysis of fMRI data from normal and clinical populations, especially for uncovering unpredictable transient patterns of brain activity associated with performance of psychomotor tasks.

Algorithms

Blind separation of auditory event-related brain responses into independent components.

Averaged event-related potential (ERP) data recorded from the human scalp reveal electroencephalographic (EEG) activity that is reliably time-locked and phase-locked to experimental events. We report here the application of a method based on information theory that decomposes one or more ERPs recorded at multiple scalp sensors into a sum of components with fixed scalp distributions and sparsely activated, maximally independent time courses. Independent component analysis (ICA) decomposes ERP data into a number of components equal to the number of sensors. The derived components have distinct but not necessarily orthogonal scalp projections. Unlike dipole-fitting methods, the algorithm does not model the locations of their generators in the head. Unlike methods that remove second-order correlations, such as principal component analysis (PCA), ICA also minimizes higher-order dependencies. Applied to detected-and undetected-target ERPs from an auditory vigilance experiment, the algorithm derived ten components that decomposed each of the major response peaks into one or more ICA components with relatively simple scalp distributions. Three of these components were active only when the subject detected the targets, three other components only when the target went undetected, and one in both cases. Three additional components accounted for the steady-state brain response to a 39-Hz background click train. Major features of the decomposition proved robust across sessions and changes in sensor number and placement. This method of ERP analysis can be used to compare responses from multiple stimuli, task conditions, and subject states.

Algorithms

Estimating alertness from the EEG power spectrum.

In tasks requiring sustained attention, human alertness varies on a minute time scale. This can have serious consequences in occupations ranging from air traffic control to monitoring of nuclear power plants. Changes in the electroencephalographic (EEG) power spectrum accompany these fluctuations in the level of alertness, as assessed by measuring simultaneous changes in EEG and performance on an auditory monitoring task. By combining power spectrum estimation, principal component analysis and artificial neural networks, we show that continuous, accurate, noninvasive, and near real-time estimation of an operator's global level of alertness is feasible using EEG measures recorded from as few as two central scalp sites. This demonstration could lead to a practical system for noninvasive monitoring of the cognitive state of human operators in attention-critical settings.

Acoustic Stimulation

Effects of voluntary movements on early auditory brain responses.

It has not been clear whether or not early information processing in the human auditory cortex is altered by voluntary movements. We report a movement-related, complex event-related potential consisting of relatively long-lasting amplitude and phase perturbations induced in an ongoing auditory steady-state response (SSR) by brief self-paced finger movements. Our results suggest that processing in the auditory cortex during the first 50-100 ms after stimulus delivery is affected before, during, and after voluntary movements, beginning with a 1- to 2-ms delay in the SSR wave form starting 1-2 s before the movement.

Acoustic Stimulation

Tonic, phasic, and transient EEG correlates of auditory awareness in drowsiness.

During drowsiness, human performance in responding to above-threshold auditory targets tends to vary irregularly over periods of 4 min and longer. These performance fluctuations are accompanied by distinct changes in the frequency spectrum of the electroencephalogram (EEG) on three time scales: (1) during minute-scale and longer periods of intermittent responding, mean activity levels in the (< 4 Hz) delta and (4-6 Hz) theta bands, and at the sleep spindle frequency (14 Hz) are higher than during alert performance. (2) In most subjects, 4-6 Hz theta EEG activity begins to increase, and gamma band activity above 35 Hz begins to decrease, about 10 s before presentations of undetected targets, while before detected targets, 4-6 Hz amplitude decreases and gamma band amplitude increases. Both these amplitude differences last 15-20 s and occur in parallel with event-related cycles in target detection probability. In the same periods, alpha and sleep-spindle frequency amplitudes also show prominent 15-20 s cycles, but these are not phase locked to performance cycles. (3) A second or longer after undetected targets, amplitude at intermediate (10-25 Hz) frequencies decreases briefly, while detected targets are followed by a transient amplitude increase in the same latency and frequency range.

Acoustic Stimulation

Changes in alertness are a principal component of variance in the EEG spectrum.

Minute-scale fluctuations in the normalized EEG log spectrum, when correlated with concurrent changes in level of performance on a sustained auditory detection task, showed that a single principal component of EEG spectral variance is linearly related to minute-scale changes in detection performance. The particular EEG frequencies at which this coupling is expressed are similar for most subjects under a range of task conditions, and match those recently reported from analysis of verbal self-reports during drowsiness. The one-dimensional relationship between detection performance and the EEG spectrum confirms quantitatively the intuitive assumption that minute-scale changes in behavioral alertness during drowsiness are predominantly linked to changes in global brain dynamics along a single dimension of psychophysiological arousal.

Acoustic Stimulation

Different event-related patterns of gamma-band power in brain waves of fast- and slow-reacting subjects.

Fast- and slow-reacting subjects exhibit different patterns of gamma-band electroencephalogram (EEG) activity when responding as quickly as possible to auditory stimuli. This result appears to confirm long-standing speculations of Wundt that fast- and slow-reacting subjects produce speeded reactions in different ways and demonstrates that analysis of event-related changes in the amplitude of EEG activity recorded from the human scalp can reveal information about event-related brain processes unavailable using event-related potential measures. Time-varying spectral power in a selected (35- to 43-Hz) gamma frequency band was averaged across trials in two experimental conditions: passive listening and speeded reacting to binaural clicks, forming 40-Hz event-related spectral responses. Factor analysis of between-subject event-related spectral response differences split subjects into two near-equal groups composed of faster- and slower-reacting subjects. In faster-reacting subjects, 40-Hz power peaked near 200 ms and 400 ms poststimulus in the react condition, whereas in slower-reacting subjects, 40-Hz power just before stimulus delivery was larger in the react condition. These group differences were preserved in separate averages of relatively long and short reaction-time epochs for each group. gamma-band (20-60 Hz)-filtered event-related potential response averages did not differ between the two groups or conditions. Because of this and because gamma-band power in the auditory event-related potential is small compared with the EEG, the observed event-related spectral response features must represent gamma-band EEG activity reliably induced by, but not phase-locked to, experimental stimuli or events.

Acoustic Stimulation

Oscillatory brain activity during a motor task.

Changes in spectral power in neuromagnetic fields associated with a manual task requiring a high level of sensorimotor integration (SMI) were investigated by analysing spontaneous, non-invasively recorded activity during motor preparation (WAIT), task performance (SMI), and control (REST) conditions in four healthy, right-handed human subjects. Neuromagnetic fields were recorded over the left sensorimotor cortex using a 37-channel instrument. In all subjects, a prominent narrow-band motor preparation rhythm centered near 19 Hz was consistently observed during the WAIT state. During SMI, mean relative increases in 26-30 Hz activity appeared in two of the subjects, paralleling gamma band enhancement recently observed during SMI in monkeys.

Adult

Lapses in alertness: coherence of fluctuations in performance and EEG spectrum.

Thirteen subjects detected noise burst targets presented in a white noise background at a mean rate of 10/min. Within each session, local error rate, defined as the fraction of targets detected in a 33 sec moving window, fluctuated widely. Mean coherence between slow mean variations in EEG power and in local error rate was computed for each EEG frequency and performance cycle length, and was shown by a Monte Carlo procedure to be significant for many EEG frequencies and performance cycle lengths, particularly in 4 well-defined EEG frequency bands, near 3, 10, 13, and 19 Hz, and at higher frequencies in two cycle length ranges, one longer than 4 min and the other near 90 sec/cycle. The coherence phase plane contained a prominent phase reversal near 6 Hz. Sorting individual spectra by local error rate confirmed the close relation between performance and EEG power and its relative within-subject stability. These results show that attempts to maintain alertness in an auditory detection task result in concurrent minute and multi-minute scale fluctuations in performance and the EEG power spectrum.

Acoustic Stimulation

Auditory event-related dynamics of the EEG spectrum and effects of exposure to tones.

A new measure of event-related brain dynamics, the event-related spectral perturbation (ERSP), is introduced to study event-related dynamics of the EEG spectrum induced by, but not phase-locked to, the onset of the auditory stimuli. The ERSP reveals aspects of event-related brain dynamics not contained in the ERP average of the same response epochs. Twenty-eight subjects participated in daily auditory evoked response experiments during a 4 day study of the effects of 24 h free-field exposure to intermittent trains of 89 dB low frequency tones. During evoked response testing, the same tones were presented through headphones in random order at 5 sec intervals. No significant changes in behavioral thresholds occurred during or after free-field exposure. ERSPs induced by target pips presented in some inter-tone intervals were larger than, but shared common features with, ERSPs induced by the tones, most prominently a ridge of augmented EEG amplitude from 11 to 18 Hz, peaking 1-1.5 sec after stimulus onset. Following 3-11 h of free-field exposure, this feature was significantly smaller in tone-induced ERSPs; target-induced ERSPs were not similarly affected. These results, therefore, document systematic effects of exposure to intermittent tones on EEG brain dynamics even in the absence of changes in auditory thresholds.

Acoustic Stimulation

Physiological studies of central masking in man. I: The effects of noise on the 40-Hz steady-state response.

In a typical masking situation, two Békésy waves overlap on the basilar membrane, and each of them initiates a stream of nerve impulses that enters the brain via the auditory nerve. Much is known about the overlapping of the cochlear waves, but much less about where, how, and even if at all, the impulse streams interact once they get inside the brain. In these experiments the incoming impulses are measured electrophysiologically using the auditory brainstem response (ABR), and, simultaneously, using the 40-Hz auditory steady-state response (SSR) to monitor events at a probable site of their interaction, the auditory cortex. The principal finding is that, when progressively increasing levels of continuous noise are presented to the contralateral ear, the SSR to the signal drops to about half its control amplitude. Second, low levels of ipsilateral noise reliably enhance SSR amplitude. Third, moderate levels of ipsilateral noise reduce SSR latency. In none of these cases does the ABR show similar effects. These findings are interpreted to mean that, in each case, impulses excited by the signal interact with impulses excited by the noise, and regardless of ear of origin the interactions take place beyond the brainstem level where ABR wave V is generated, either before the impulses reach the cortex, or in the cortex itself.

Acoustic Stimulation

Physiological studies of central masking in man. II: Tonepip SSRs and the masking level difference.

The auditory steady-state response (SSR), an evoked response generated in the auditory cortex, was initiated by monaural trains of 500-Hz tonepips repeated at rates near 40 Hz while wideband noise was being delivered to the same or opposite ear. Contralateral noise reduced SSR amplitudes in an intensity-dependent manner, whereas ipsilateral noise enhanced the SSR amplitudes at low levels and depressed them at high levels. Systematic phase changes accompanied the amplitude changes. These results, obtained with tonepips, closely resemble those previously reported for clicks. A third experiment, a masking level difference (MLD) experiment, examined changes in the SSR measures during four successive tonepip-plus-noise conditions: (1) monaural tonepips alone; (2) adding ipsilateral noise; (3) then adding contralateral noise; (4) finally, adding contralateral tonepips. The SSR amplitude changes measured in the experiment did not always correspond with the changes in perception reported by the subject.

Acoustic Stimulation

Human auditory evoked gamma-band magnetic fields.

We have discovered a ca. 40-Hz transient magnetic oscillatory response, evoked in the human brain by the onset of auditory stimuli, consisting of four or more cycles locked in phase to stimulus onset in approximately the 20- to 130-ms poststimulus interval. The response originates in the supratemporal auditory cortex, some millimeters deeper and anterior to the source of the larger-amplitude slow-wave M100 component of the evoked magnetic field and moves in a posterior arcing trajectory 1 cm or more in length. The oscillatory cortical activation elicited by auditory stimuli may be similar to the gamma-band cortical oscillations elicited by olfactory and visual stimuli and may represent an essential component of auditory perceptual processing.

Acoustic Stimulation

Auditory steady-state responses: threshold prediction using phase coherence.

These experiments add a measure of response phase variance--'phase coherence'--to the analysis procedures applied to auditory steady-state responses (SSR). The effects of stimulus frequency, intensity, rate and total number (i.e., recording time) were studied using 11 normal adult subjects. In a first experiment, SSR phase coherence was found to be highest at presentation rates near 40/sec, even when response amplitudes were higher at other rates. Further, phase coherence was observed to be linearly related (r = 0.91) to signal-to-noise ratio. Two further experiments demonstrated that phase coherence can correctly detect responses to near-threshold stimuli. In 15 min runs, significant phase coherence was detected within 6 dB of behavioral threshold in 6 subjects for 0.5 and 2.0 kHz signals, while phase coherence in no-stimulus control runs did not reach significance. Minimum data collection time required to record significant (P less than 0.01) responses was studied for 10 subjects. In 2 of 40 recordings at 10 dB SL phase coherence remained insignificant after even 15 min. However, average recording time to reach significance at 10 dB SL was less than 4 min in 38 of 40 recordings, and less than 1 min at 25 dB SL (18 of 18 recordings). These results indicate that using phase coherence to detect the presence of the 40/sec auditory steady-state response, efficient threshold search procedures may be devised to provide fast, accurate, and objective estimates of auditory behavioral thresholds in nearly all normal adults.

Acoustic Stimulation

A 40-Hz auditory potential recorded from the human scalp.

Computer techniques readily extract from the brainwaves an orderly sequence of brain potentials locked in time to sound stimuli. The potentials that appear 8 to 80 msec after the stimulus resemble 3 or 4 cycles of a 40-Hz sine wave; we show here that these waves combined to form a single, stable, composite wave when the sounds are repeated at rates around 40 per sec. This phenomenon, the 40-Hz event-related potential (ERP), displays several properties of theoretical and practical interest. First, it reportedly disappears with surgical anesthesia, and it resembles similar phenomena in the visual and olfactory system, facts which suggest that adequate processing of sensory information may require cyclical brain events in the 30- to 50-Hz range. Second, latency and amplitude measurements on the 40-Hz ERP indicate it may contain useful information on the number and basilar membrane location of the auditory nerve fibers a given tone excites. Third, the response is present at sound intensities very close to normal adult thresholds for the audiometric frequencies, a fact that could have application in clinical hearing testing.

Auditory Pathways