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Electroencephalographic measurement of motor cortex control of muscle activity in humans.

OBJECTIVE: To detect and measure correlation between cortical and muscle activities, coherence analysis was used. METHODS: The electroencephalogram (EEG) and electromyogram (EMG) were recorded in 9 normal volunteers during tonic contraction of upper and lower limb muscles on the right side. Coherence between EEG and EMG was computed to analyze their linear association. RESULTS: EEG over the contralateral sensorimotor area was coherent with EMG, with peak coherence at 11-36 Hz (mean, 22 Hz). For the abductor pollicis brevis (APB) muscle, peak coherence, as determined by functional brain mapping with focal transcranial magnetic stimulation (TMS), was over or slightly posterior to the hand area on the primary motor cortex determined by focal transcranial magnetic stimulation (TMS). Peak coherence over the scalp was somatotopically organized. The temporal relation between EEG and EMG was analyzed with a new model for interpreting the phase shift ('constant phase shift plus constant time lag' model). For the APB muscle, the phase relation between cortical and muscular oscillations differed in the frequency ranges of 3-13 Hz and 14-50 Hz, respectively, suggesting that different coupling mechanisms operate in different bands. Only the phase shift between cortical and motoneuronal firing at 14-50 Hz was reliably estimated by a linear model. At 14-50 Hz, motoneuronal firing was led by surface-negative cortical activity with a constant time lag that depended on the cortical-muscular distance. For the APB muscle, the time lag was slightly shorter than the cortical-muscular conduction time determined by TMS. Vibratory stimulation (100 Hz) of a muscle tendon during tonic contraction had no significant effect on cortical-muscular coherence, indicating that cortical oscillation reflected motor rather than sensory activity. CONCLUSIONS: The present findings suggest temporal coding of the oscillatory motor control system (3-13 Hz vs. 14-50 Hz), and confirm the functional importance of cortical beta and gamma rhythms in the motor efferent command. Cortical-muscular synchronization is most likely mediated by the direct corticospinal pathway within the frequency range of 14-50 Hz.

Adult↗

Brain-computer communication based on the dynamics of brain oscillations.

This chapter presents a review of brain-computer communication based on motor imagery and the dynamics of brain oscillations. The concept of motor imagery as experimental strategy and the two different modes of operation a brain-computer interface can have are explained. An EEG based brain switch that can control a FES-induced hand grasp of a tetraplegic and an approach towards an ECoG based brain switch are presented.

Brain↗

ERD as an index of anticipatory attention? Effects of stimulus degradation.

Previous research has suggested that the stimulus-preceding negativity (SPN) is largely independent of stimulus modality. In contrast, the scalp topography of the event-related desynchronization (ERD) related to the anticipation of stimuli providing knowledge of results (KR) is modality dependent. These findings, combined with functional SPN research, lead to the hypothesis that anticipatory ERD reflects anticipatory attention, whereas the SPN mainly depends on the affective-motivational properties of the anticipated stimulus. To further investigate the prestimulus ERD, and compare this measure with the SPN, 12 participants performed a time-estimation task, and were informed about the quality of their time estimation by an auditory or a visual stimulus providing KR. The KR stimuli could be either intact or degraded. Auditory degraded KR stimuli were less effective than other KR stimuli in guiding subsequent behavior, and were preceded by a larger SPN. There were no effects of degradation on the SPN in the visual modality. Preceding auditory KR stimuli no ERD was present, whereas preceding visual stimuli an occipital ERD was found. However, contrary to expectation, the latter was larger preceding intact than preceding degraded stimuli. It is concluded that the data largely agree with an interpretation of the pre-KR SPN as a reflection of the anticipation of the affective-motivational value of KR stimuli, and of the prestimulus ERD as a perceptual anticipatory attention process.

Acoustic Stimulation↗

Working memory retention systems: a state of activated long-term memory.

High temporal resolution event-related brain potential and electroencephalographic coherence studies of the neural substrate of short-term storage in working memory indicate that the sustained coactivation of both prefrontal cortex and the posterior cortical systems that participate in the initial perception and comprehension of the retained information are involved in its storage. These studies further show that short-term storage mechanisms involve an increase in neural synchrony between prefrontal cortex and posterior cortex and the enhanced activation of long-term memory representations of material held in short-term memory. This activation begins during the encoding/comprehension phase and evidently is prolonged into the retention phase by attentional drive from prefrontal cortex control systems. A parsimonious interpretation of these findings is that the long-term memory systems associated with the posterior cortical processors provide the necessary representational basis for working memory, with the property of short-term memory decay being primarily due to the posterior system. In this view, there is no reason to posit specialized neural systems whose functions are limited to those of short-term storage buffers. Prefrontal cortex provides the attentional pointer system for maintaining activation in the appropriate posterior processing systems. Short-term memory capacity and phenomena such as displacement of information in short-term memory are determined by limitations on the number of pointers that can be sustained by the prefrontal control systems.

Animals↗

Hallucinations in schizophrenia, sensory impairment, and brain disease: a unifying model.

Based on recent insight into the thalamocortical system and its role in perception and conscious experience, a unified pathophysiological framework for hallucinations in neurological and psychiatric conditions is proposed, which integrates previously unrelated neurobiological and psychological findings. Gamma-frequency rhythms of discharge activity from thalamic and cortical neurons are facilitated by cholinergic arousal and resonate in networks of thalamocortical circuits, thereby transiently forming assemblies of coherent gamma oscillations under constraints of afferent sensory input and prefrontal attentional mechanisms. If perception is based on synchronisation of intrinsic gamma activity in the thalamocortical system, then sensory input to specific thalamic nuclei may merely play a constraining role. Hallucinations can be regarded as underconstrained perceptions that arise when the impact of sensory input on activation of thalamocortical circuits and synchronisation of thalamocortical gamma activity is reduced. In conditions that are accompanied by hallucinations, factors such as cortical hyperexcitability, cortical attentional mechanisms, hyperarousal, increased noise in specific thalamic nuclei, and random sensory input to specific thalamic nuclei may, to a varying degree, contribute to underconstrained activation of thalamocortical circuits. The reticular thalamic nucleus plays an important role in suppressing random activity of relay cells in specific thalamic nuclei, and its dysfunction may be implicated in the biological vulnerability to hallucinations in schizophrenia. Combined with general activation during cholinergic arousal, this leads to excessive disinhibition in specific thalamic nuclei, which may allow cortical attentional mechanisms to recruit thalamic relay cells into resonant assemblies of gamma oscillations, regardless of their actual sensory input, thereby producing an underconstrained perceptual experience.

Arousal↗

An evolutionary theory of schizophrenia: cortical connectivity, metarepresentation, and the social brain.

Schizophrenia is a worldwide, prevalent disorder with a multifactorial but highly genetic aetiology. A constant prevalence rate in the face of reduced fecundity has caused some to argue that an evolutionary advantage exists in unaffected relatives. Here, I critique this adaptationist approach, and review--and find wanting--Crow's "speciation" hypothesis. In keeping with available biological and psychological evidence, I propose an alternative theory of the origins of this disorder. Schizophrenia is a disorder of the social brain, and it exists as a costly trade-off in the evolution of complex social cognition. Paleoanthropological and comparative primate research suggests that hominids evolved complex cortical interconnectivity (in particular, frontotemporal and frontoparietal circuits) to regulate social cognition and the intellectual demands of group living. I suggest that the ontogenetic mechanism underlying this cerebral adaptation was sequential hypermorphosis and that it rendered the hominid brain vulnerable to genetic and environmental insults. I argue that changes in genes regulating the timing of neurodevelopment occurred prior to the migration of Homo sapiens out of Africa 100,000-150,000 years ago, giving rise to the schizotypal spectrum. While some individuals within this spectrum may have exhibited unusual creativity and iconoclasm, this phenotype was not necessarily adaptive in reproductive terms. However, because the disorder shared a common genetic basis with the evolving circuitry of the social brain, it persisted. Thus schizophrenia emerged as a costly trade-off in the evolution of complex social cognition.

Animals↗

Cortical mapping of gamma oscillations in areas V1 and V4 of the macaque monkey.

To characterize the temporal and spatial parameters of gamma activity evoked by visual stimuli in areas V1 and V4 of the monkey cortex, we recorded the electrocorticogram (ECoG) with an implanted array of 28 and 31 subdural electrodes placed over the surface of the operculum in two anesthetized monkeys. This intermediate level of recordings should help to bridge the gap between multiunit and scalp recordings. Both averaged and single-trial responses to small flashed stimuli, for which we varied the retinotopic position, the luminance and the color, were analyzed in the time-frequency domain using a wavelet-based decomposition of the signal. Large gamma oscillations (40-55 Hz), not phase locked to stimulus onset, were observed during the whole stimulus presentation, whereas visual evoked potentials (VEPs) were present mainly at stimulus onset and offset. Cortical mapping showed that both activities were restricted in spatial extent and followed the retinotopic organization of area V1 on the operculum, thus strongly suggesting they were generated in the underlying cortex. Oscillatory burst detection in single trials showed that one to two bursts lasting from 100 ms to 500 ms occurred in the first 500 ms following stimulus onset, and that bursts occurring during the subsequent phases of the response had a smaller amplitude and duration. Finally, we showed that gamma activity was stronger with higher luminances and for red than for green, yellow, or white stimuli of same luminance. In one animal we recorded gamma activity over area V4. This was of lower magnitude than the activity recorded over V1 and was delayed by 40 ms with respect to the beginning of gamma activity in V1, in contrast with the VEPs that were delayed by 20 ms only. Both gamma oscillations and early VEP followed the retinotopic organization of V4 over the prelunate gyrus. The results show that gamma oscillations are dependent upon the same parameters as the VEPs (retinotopic position, luminance, and color). However, the differences in the time course of VEPs and gamma activity (transient vs. sustained) suggests that these two responses may reflect different cell populations, different networks, or different firing modes.

Animals↗

Long-range interactions modulate the contrast gain in the lateral geniculate nucleus of cats.

In previous work, we have shown that sudden image displacements well outside the classical receptive field modulate the visual sensitivity of LGN relay cells. Here we report the effect of image displacements on the response versus contrast function. The stimuli consisted of a central spot of optimal size and polarity (contrast range: 3-98%), flashed alone or in the presence of a peripheral annulus (radii: 5-15 deg) containing a low spatial-frequency grating displaced at saccade-like velocities (shift). The most consistent effect of the shift on the response to a central spot was to reduce the responsiveness of Y relay cells and, to a lesser extent, of X relay cells. The reduction in responsiveness was primarily a divisive rather than a subtractive effect and could be modelled by assuming that a greater contrast was required to produce a given excitatory response. In the absence of a central spot, remote motion had inhibitory effects on the firing rates of the majority of relay cells, but its effect on retinal ganglion cells was mainly excitatory. When the shifting grating covered the classical receptive field and its periphery, facilitatory effects or suppressive effects, depending on the spatial phase of the pattern, were observed in both retinal and geniculate cells. Remote motion strongly suppresses the responsiveness of relay cells to stimuli within the classical receptive field. This suppressive effect involves intrageniculate processing and is primarily associated with a reduction in contrast gain. It is likely that shift suppression contributes to the loss of visual sensitivity observed in saccadic suppression.

Animals↗

Emergent oscillations in a realistic network: the role of inhibition and the effect of the spatiotemporal distribution of the input.

We have simulated a network of 10,000 two-compartment cells, spatially distributed on a two-dimensional sheet; 15% of the cells were inhibitory. The input to the network was spatially delimited. Global oscillations frequently were achieved with a simple set of connectivity rules. The inhibitory neurons paced the network, whereas the excitatory neurons amplified the input, permitting oscillations at low-input intensities. Inhibitory neurons were active over a greater area than excitatory ones, forming a ring of inhibition. The oscillation frequency was modulated to some extent by the input intensity, as has been shown experimentally in the striate cortex, but predominantly by the properties of the inhibitory neurons and their connections: the membrane and synaptic time constants and the distribution of delays. In networks that showed oscillations and in those that did not, widely distributed inputs could lead to the specific recruitment of the inhibitory neurons and to near zero activity of the excitatory cells. Hence the spatial distribution of excitatory inputs could provide a means of selectively exciting or inhibiting a target network. Finally, neither the presence of oscillations nor the global spike activity provided any reliable indication of the level of excitatory output from the network.

Computer Systems↗

Turning on and off with excitation: the role of spike-timing asynchrony and synchrony in sustained neural activity.

Delay-related sustained activity in the prefrontal cortex of primates, a neurological analogue of working memory, has been proposed to arise from synaptic interactions in local cortical circuits. The implication is that memories are coded by spatially localized foci of sustained activity. We investigate the mechanisms by which sustained foci are initiated, maintained, and extinguished by excitation in networks of Hodgkin-Huxley neurons coupled with biophysical spatially structured synaptic connections. For networks with a balance between excitation and inhibition, a localized transient stimulus robustly initiates a localized focus of activity. The activity is then maintained by recurrent excitatory AMPA-like synapses. We find that to maintain the focus, the firing must be asynchronous. Consequently, inducing transient synchrony through an excitatory stimulus extinguishes the sustained activity. Such a monosynaptic excitatory turn-off mechanism is compatible with the working memory being wiped clean by an efferent copy of the motor command. The activity that codes working memories may be structured so that the motor command is both the read-out and a direct clearing signal. We show examples of data that is compatible with our theory.

Action Potentials↗

Effects of amitriptyline on diurnal variations of oxidation-reduction enzyme activities in rat lymphocytes during experimental desynchronosis.

We studied the effects of amitriptyline on diurnal variations of oxidative-reduction enzyme activities (succinate dehydrogenase, lactate dehydrogenase, and NADPH diaphorase) in rat peripheral blood lymphocytes during experimental desynchronosis. Desynchronosis was induced by constant light exposure for 14 days and manifested in a morning shift of maximum lactate dehydrogenase activity and elevated morning NADPH diaphorase activity. Amitriptyline normalized morning lactate dehydrogenase activity and diurnal variations in NADPH diaphorase activity.

Amitriptyline↗

Genetic and environmental influences on EEG coherence.

EEG coherence measures the covariation in electrical brain activity between two locations on the scalp and is used to study connectivity between cortical regions. The aim of this study was to determine the heritability of EEG coherence. Coherence was measured in a group of 213 16-yr-old twin pairs. By including male and female twin pairs in the sample, sex differences in genetic architecture were systematically examined. The EEG was obtained during quiet supine resting. Coherence was estimated for short and long distance combinations of electrode pairs along the anterior-posterior axis within a hemisphere for four frequency bands (delta, theta, alpha and beta). Averaged over all electrode combinations about 60% of the variance was explained by genetic factors for coherence in the theta, alpha and beta bands. For the delta band, the heritability was somewhat lower. No systematic sex differences in genetic architecture were found. All environmental influences were nonshared, i.e., unique factors including measurement error. Environmental factors shared by twin siblings did not influence variation in EEG coherence. These results suggest that individual differences in coherence form a potential candidate for (molecular) genetic studies on brain function.

Adolescent↗

The functional significance of absolute power with respect to event-related desynchronization.

The question is examined whether the extent of changes in relative band power as measured by event-related desynchronization (ERD) depends on absolute band power. The results for target stimuli of a simple oddball task indicate that the prestimulus (reference) level of absolute band power has indeed a strong influence on ERD. Whereas for the alpha band large band power in the reference interval is related to a strong degree of alpha suppression as measured by ERD, the opposite holds true for the theta band. Here, a low level of band power during the reference interval is related to a pronounced increase in band power during the processing of the target stimulus. In contrast to alpha and theta, ERD in the delta band is not influenced by the magnitude of band power in the reference interval.

Adolescent↗

Spatial EEG synchronisation over sensorimotor hand areas in brisk and slow self-paced index finger movements.

The changes of spatial EEG synchronisation during brisk and slow voluntary self-paced movements of the right and left index finger were analysed in 12 right-handed and 11 left-handed subjects. EEG was recorded from the left and right sensorimotor area using 24 closely spaced electrodes. A novel measure of spatial EEG synchronisation, omega-complexity, was computed separately for the left and right sensorimotor area in 64 overlapping one-second epochs representing 4.5 s of the pre-movement and 3.5 s of the post-movement period. Omega-complexity was higher, hence spatial synchronisation was lower, in slow than in brisk movements, especially in the right-handed. A sustained increase of omega-complexity was observed during execution of a slow movement. A decrease of omega-complexity which was often associated with a brief burst of spatially synchronised 10-Hz oscillations occurred at the onset of extensor muscle contraction. We suggest that increased spatial EEG synchronisation at movement onset may prevent "spillover" of excitation from the sensorimotor hand area to other cortical regions. During movement, the cortical neuronal assemblies subserve distinct, specialised functions manifesting in increased omega-complexity.

Adult↗

New methods for three dimensional mapping of brain waves.

The representation of spatiotemporal pattern of brain electrical activity is an important issue in the research of neural assemblies. This paper describes methods of parametric 3D head or brain modeling and its corresponding interpolation for functional imaging based on brain waves. The 3D interpolation method is an extension of cortical imaging technique. It can be used with features of either transformed domains or higher order statistics of brain waves on realistic head or brain models. The simulating result suggests that it is a better method in comparison with the global nearest neighbor technique. With those techniques, an example of brainwave imaging was given using partial coherence of event-related potentials estimated by multivariate autoregressive model.

Brain Mapping↗

Left frontal EEG coherence reflects modality independent language processes.

Previous studies showed that distinct components of higher cognitive processes like memorizing of words could be correlated with changes in different frequency bands of the human EEG. This study was designed in order to find out if 1) some frequency bands show power and coherence changes only due to the modality of presented stimuli (either auditory or visual) and 2) if other frequency bands show modality independent effects which should reflect real cognitive-linguistic differences between word classes (either concrete and abstract nouns). EEG was recorded from sixteen right-handed females which had to memorize auditorily and visually presented concrete and abstract nouns. Results show the alpha-1 band to reveal no differences between word classes but demonstrate an influence of modality of stimulus presentation. The only modality independent differences between concrete and abstract noun processing were found in the delta, theta and beta-1 band at left frontal electrodes.

Alpha Rhythm↗

Tuning neocortical pyramidal neurons between integrators and coincidence detectors.

Do cortical neurons operate as integrators or as coincidence detectors? Despite the importance of this question, no definite answer has been given yet, because each of these two views can find its own experimental support. Here we investigated this question using models of morphologically-reconstructed neocortical pyramidal neurons under in vivo like conditions. In agreement with experiments we find that the cell is capable of operating in a continuum between coincidence detection and temporal integration, depending on the characteristics of the synaptic inputs. Moreover, the presence of synaptic background activity at a level comparable to intracellular measurements in vivo can modulate the operating mode of the cell, and act as a switch between temporal integration and coincidence detection. These results suggest that background activity can be viewed as an important determinant of the integrative mode of pyramidal neurons. Thus, background activity not only sharpens cortical responses but it can also be used to tune an entire network between integration and coincidence detection modes.

Action Potentials↗