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Small-world network organization of functional connectivity of EEG slow-wave activity during sleep.

OBJECTIVE: To analyze the functional connectivity patterns of the EEG slow-wave activity during the different sleep stages and Cyclic Alternating Pattern (CAP) conditions, using concepts derived from Graph Theory. METHODS: We evaluated spatial patterns of EEG slow-wave synchronization between all possible pairs of electrodes (19) placed over the scalp of 10 sleeping healthy young normal subjects using two graph theoretical measures: the clustering coefficient (Cp) and the characteristic path length (Lp). The measures were obtained during the different sleep stages and CAP conditions from the real EEG connectivity networks and randomized control (surrogate) networks (Cp-s and Lp-s). RESULTS: Cp and Cp/Cp-s increased significantly from wakefulness to sleep while Lp and Lp/Lp-s did not show changes. Cp/Cp-s was higher for A1 phases, compared to B phases of CAP. CONCLUSIONS: The network organization of the EEG slow-wave synchronization during sleep shows features characteristic of small-world networks (high Cp combined with low Lp); this type of organization is slightly but significantly more evident during the CAP A1 subtypes. SIGNIFICANCE: Our results show feasibility of using graph theoretical measures to characterize the complexity of brain networks during sleep and might indicate sleep, and the A1 phases of CAP in particular, as a period during which slow-wave synchronization shows optimal network organization for information processing.

Adult↗

Prestimulus EEG alpha phase synchronicity influences N100 amplitude and reaction time.

The influence of the level of prestimulus alpha phase synchronicity on the N100 component and reaction time (RT) was examined, in target auditory oddball data from 25 normal subjects. Alpha phase synchronicity is a new measure consisting (for a given stimulus presentation) of the angular or circular variance of the alpha phase at stimulus onset across the parieto-occipital sites. The lower the angular variance, the higher the phase synchronicity (the more closely in phase the alpha activity across these sites) and vice versa. Subaveraged event-related potentials (ERPs) were formed for high and low prestimulus alpha phase synchronicity stimulus presentations. N100 amplitude was significantly greater in the high than the low phase synchronicity subaverages. In addition, RT was significantly reduced in the high prestimulus alpha phase synchronicity cases. Alpha phase synchronicity reflects an aspect of brain state that influences subsequent stimulus processing.

Adult↗

Chronic pretrigeminal and cerveau isolé cats.

Ten pretrigeminal and ten cerveau isole cats were observed chronically. During 24-36 h sessions EEG activity was continuously recorded and the EEG and ocular responses to visual and olfactory stimuli were studied. In the pretrigeminal cat acute and chronic stages were distinguished, and in the cerveau isole, acute, "early chronic" and "late chronic" stages. During the acute stage, the pretrigeminal cat is continuously awake, whereas the cerveau isole is comatose. During the "early chronic stage", which lasted at least about 3 weeks, the cerveau isole is semicomatose. During the chronic stage in the pretrigeminal cat and the "late chronic stage" in the cerveau isole, the sleep-waking cycle is present. In both preparations alert wakefulness, drowsiness, light .synchronized sleep and deep synchronized sleep occupy, respectively, about 30 percent, 45 percent, 15 percent and 10 percent of the time. Thus, synchronized sleep is strikingly reduced in comparison with an intact cat, while desynchronized sleep is absent.

Animals↗

Synchronization of local neural networks in the somatosensory cortex: A comparison of stationary and moving stimuli.

Spontaneous and stimulus-induced responses were recorded from neighboring groups of neurons by an array of electrodes in the primary (SI) somatosensory cortex of intact, halothane-anesthetized cats. Cross-correlation analysis was used to characterize the coordination of spontaneous activity and the responses to peripheral stimulation with moving or stationary air jets. Although synchronization was detected in only 10% (88 of 880) of the pairs of single neurons that were recorded, cross-correlation analysis of multiunit responses revealed significant levels of synchronization in 64% of the 123 recorded electrode pairs. Compared with spontaneous activity, both stationary and moving air jets caused substantial increases in the rate, proportion, and temporal precision of synchronized activity in local regions of SI cortex. Among populations of neurons that were synchronized by both types of air-jet stimulation, the mean rate of synchronized activity was significantly higher during moving air-jet stimulation than during stationary air-jet stimulation. Moving air jets also produced significantly higher correlation coefficients than stationary air jets in the raw cross-correlograms (CCGs) but not in the shift-corrected CCGs. The incidence and rate of stimulus-induced synchronization varied with the distance separating the recording sites. For sites separated by </=300 microm, 80% of the multiunit responses displayed significant levels of synchronization during both types of air-jet stimulation. For sites separated by >/=500 microm, only 37% of the multiunit responses were synchronized by discrete stimulation with a single air jet. Measurements of the multiunit CCG peak half-widths showed that the correlated activity produced by moving air jets had slightly less temporal variability than that produced by stationary air jets. These results indicate that moving stimuli produce greater levels of synchronization than stationary stimuli among local groups of SI neurons and suggest that neuronal synchronization may supplement the changes in firing rate which code intensity and other attributes of a cutaneous stimulus.

Animals↗

Flash-related synchronization and desynchronization revealed by a multiple band frequency analysis.

The fast Fourier transform (FFT) is a good method to estimate power spectral density (PSD), but the frequency resolution is limited to the sampling window, and thus the precise characteristics of PSD for short signals are not clear. To relax the limitation, a multiple band-pass filter was introduced to estimate the precise course of PSDs for flash visual evoked potentials (VEPs). Signals were recorded during -200 and 600 ms using balanced noncephalic electrodes, and sampled at 1,000 Hz in 12 bits. With 1 Hz and 10 ms resolutions, PSDs were estimated between 10 and 100 Hz. Background powers at the alpha- and beta-bands were high over the posterior scalp, and powers around 200 ms were evoked at the same bands over the same region, corresponding to P110 and N165 of VEPs. Normalized PSDs showed evoked powers around 200 ms and suppressed powers following the evoked powers over the posterior scalp. The evoked powers above the 20 Hz band were not statistically significant, however, the gamma band was significantly evoked intra-individually; details in the gamma bands were varied among the subjects. Details of PSDs were complicated even for a simple task such as watching flashes; both synchronization and desynchronization occurred with different distributions and different time courses.

Adult↗

[The mechanisms of the formation and the role of the oscillatory activity of the neuronal populations in brain systemic activities].

Experimental findings and theoretical concepts that have led to a new insight into the neurophysiological mechanisms underlying information processing and brain state are presented in this review. It is assumed that the brain information processing associated with elementary neuronal assembles is reflected by oscillations in the range of 30-70 Hz and their spatio-temporal organization in millisecond intervals. He functional brain state associated with the non-specific systems is reflected in synchronization or desynchronization of activity of large neuronal populations in the frequency range of 20 Hz.

Brain↗

The role of sensory information in the production of periodic finger-tapping sequences.

A subject lacking proprioceptive and tactile sensibility below the neck and a group of control subjects performed sequences of periodic finger taps involving a pattern of accentuation. The required intertap interval was 700 ms. In some situations, the taps were synchronized with the clicks of a metronome. Feedback conditions were manipulated by either allowing or not allowing the subjects to hear the taps and see their finger movements. We recorded the trajectory of the subjects' finger displacement in the vertical plane, and the force and moment of contact of the finger with the response key. The control subjects achieved precise timing of the finger taps by trading off downstroke onset for movement duration, e.g., they initiated shorter-duration tapping movements with a delay. This strategy did not vary depending on task demands (e.g., synchronization) or feedback conditions. The deafferented patient produced intertap intervals on average close to the required value. However, his tap timing was characterized by increased variability and severe distortion (lengthening) after the accentuated tap, regardless of feedback conditions. He did not manifest the compensatory strategy whereby, in control subjects, movement onset was adjusted to movement duration. Thus, such a strategy in controls seems to depend on intact proprioceptive and/or tactile information from the moving limb. Upon withdrawal of visual and acoustic feedback, the deafferented subject increased the force of the taps and the amplitude of tapping movements; his mean synchronization error with the metronome also increased. However, he did not lose correct phasing between the taps and the clicks of the metronome. These findings suggest that, under normal circumstances, sequential movements are timed by an internal timekeeper which paces sensory consequences relating to the occurrence of behaviorally important events (e.g., finger taps), and not the onset of the movements eliciting those events. In the synchronization task, the timekeeper may be phase locked to the periodic acoustic stimuli by direct entrainment. Feedback information may be needed, however, for keeping any synchronization error as small as possible.

Acoustic Stimulation↗

Mechanisms to synchronize neuronal activity.

Temporal aspects of neuronal activity have received increasing attention in recent years. Oscillatory dynamics and the synchronization of neuronal activity are hypothesized to be of functional relevance to information processing in the brain. Here we review theoretical studies of single neurons at different levels of abstraction, with an emphasis on the implications for properties of networks composed of such units. We then discuss the influence of different types of couplings and choices of parameters to the existence of a stable state of synchronous or oscillatory activity. Finally we relate these theoretical studies to the available experimental data, and suggest future lines of research.

Cortical Synchronization↗

Visually evoked phase synchronization changes of alpha rhythm in migraine: correlations with clinical features.

OBJECTIVE: This study aimed to compute phase synchronization of the alpha band from a multichannel electroencephalogram (EEG) recorded under repetitive flash stimulation from migraine patients without aura. This allowed examination of ongoing EEG activity during visual stimulation in the pain-free phase of migraine. METHODS: Flash stimuli at frequencies of 3, 6, 9, 12, 15, 18, 21, 24, and 27 Hz were delivered to 15 migraine patients without aura and 15 controls, with the EEG recorded from 18 scalp electrodes, referred to the linked earlobes. The EEG signals were filtered in the alpha (7.5-13 Hz) band. For all stimulus frequencies that we evaluated, the phase synchronization index was based on the Hilbert transformation. RESULTS: Phase synchronization separated the patients and controls for the 9, 24 and 27 Hz stimulus frequencies; hyper phase synchronization was observed in patients, whereas healthy subjects were characterized by a reduced phase synchronization. These differences were found in all regions of the scalp. CONCLUSIONS: During migraine, the brain synchronizes to the idling rhythm of the visual areas under certain photic stimulations; in normal subjects however, brain regions involved in the processing of sensory information demonstrate desynchronized activity. Hypersynchronization of the alpha rhythm may suggest a state of cortical hypoexcitability during the interictal phase of migraine. SIGNIFICANCE: The employment of non-linear EEG analysis may identify subtle functional changes in the migraine brain.

Adult↗

Wavelet entropy: a new tool for analysis of short duration brain electrical signals.

Since traditional electrical brain signal analysis is mostly qualitative, the development of new quantitative methods is crucial for restricting the subjectivity in the study of brain signals. These methods are particularly fruitful when they are strongly correlated with intuitive physical concepts that allow a better understanding of brain dynamics. Here, new method based on orthogonal discrete wavelet transform (ODWT) is applied. It takes as a basic element the ODWT of the EEG signal, and defines the relative wavelet energy, the wavelet entropy (WE) and the relative wavelet entropy (RWE). The relative wavelet energy provides information about the relative energy associated with different frequency bands present in the EEG and their corresponding degree of importance. The WE carries information about the degree of order/disorder associated with a multi-frequency signal response, and the RWE measures the degree of similarity between different segments of the signal. In addition, the time evolution of the WE is calculated to give information about the dynamics in the EEG records. Within this framework, the major objective of the present work was to characterize in a quantitative way functional dynamics of order/disorder microstates in short duration EEG signals. For that aim, spontaneous EEG signals under different physiological conditions were analyzed. Further, specific quantifiers were derived to characterize how stimulus affects electrical events in terms of frequency synchronization (tuning) in the event related potentials.

Adult↗

Event-related alpha oscillations in task processing.

OBJECTIVES: Recent findings substantiate the view that electroencephalographic (EEG) alpha rhythm (7-13 Hz) is functionally involved in information processing. However, the association of alpha rhythms with cognitive brain processes is less well understood because both augmentation and suppression of alpha oscillations have been observed to accompany task performance. The present study evaluates the effect of task processing on event-related alpha oscillations at the level of single-sweep analysis. METHODS: EEG was recorded from Fz, Cz and Pz electrodes in 10 subjects participating in two experimental sessions, in which auditory stimuli with equal physical parameters were presented under different instructions (passive and task). Separate measurements of single-sweep amplitude and phase-locking were performed and statistically analyzed for consecutive time windows in the poststimulus epoch. RESULTS: Major results show that, during the cognitive task, the phase-locking of alpha oscillations at the frontal site is significantly increased for the time window of 500-1000 ms after stimulation. CONCLUSIONS: The involvement of enhanced and synchronized frontal alpha activity in higher brain processes is strongly emphasized.

Acoustic Stimulation↗

A negative component on event related potential recorded in the drowsy state.

Behavior of event related potential (ERP) components in the drowsy state was examined in nine subjects using oddball paradigm. A component with peak latency of 300 msec, N300, was superimposed on ERP in the drowsy state. N300 appeared also in stage 1 of NREM sleep and closely resembled vertex sharp wave evoked by sound stimulation in both scalp distribution and peak latency. It was suggested that N300 recorded in the drowsy state and vertex sharp wave recorded in stage 1 of NREM sleep are generated by the identical synchronizing mechanism in the brain.

Adult↗

Synchronization between background activity and visually evoked potential is not mirrored by focal hyperoxygenation: implications for the interpretation of vascular brain imaging.

We performed an electroencephalography and optical topography study simultaneously exploring electrophysiological and vascular response magnitude as a function of stimulus frequency. To elicit a response in the visual cortex, subjects were exposed to flicker frequencies varying from 1 to 25 Hz (1 Hz steps, eyes closed). Extending the standard view to compare magnitudes of the evoked neuronal to the evoked vascular response, we additionally investigated modulations of alpha-power, a marker of "background" EEG activity. The results show two discrepancies between the electrophysiological and vascular response: (1) VEP and alpha-power exhibit a discontinuous peak when stimulating at the individual alpha-frequency (IAF) (approximately 10-11 Hz), indicating resonance between background oscillations and evoked response; this is not mirrored by the vascular response. (2) The vascular response, in contrast, steadily increases up to a maximum at 7-8 Hz and slightly decreases with higher frequencies. This continuous frequency dependence is partly reflected by the decrease in alpha-power up to frequencies of 8-9 Hz and a slight increase in alpha-power beyond the IAF resonance. Although indicating an inverse relationship between alpha-power and vascular response, the frequency dependence of the evoked response does not show such a correlation. Thus, electrophysiological resonance between an individual's alpha-frequency and isofrequent stimulation is not mirrored by the vascular response. Also, spontaneous background EEG activity is an important modulator of the vascular response magnitude. We discuss these deviations from a simple one-to-one translation between evoked potential and vascular response amplitude in the light of questions concerning synchronization, attenuation, and induction of background oscillations such as the alpha-rhythm.

Adolescent↗

Influence of rostral and caudal brain stem reticular formation on thalamic neurons.

Single neuronal activity was recorded from the diffuse thalamic system. Influence of the rostral desynchronizing and caudal synchronizing structures of the brain stem reticular formation on these neurons was studied. Rostral stimulation produced an increase and caudal stimulation a decrease in the thalamic unit firing. A possible mechanism by which the brain stem reticular structures influence the cortical neurons is proposed on the basis of these findings.

Animals↗

Preictal state identification by synchronization changes in long-term intracranial EEG recordings.

OBJECTIVE: There is accumulated evidence that mesial temporal lobe seizures are preceded by a preictal transition that evolves over minutes to hours. In the present study, we investigated these possible preictal changes in long-term intracranial recordings of five patients by a measure of phase synchronization. In order to clearly distinguish preictal changes from all the other interictal states, we developed an automatic extraction of representative patterns of interictal synchronization activity. This reference library was used to classify the successive synchronization patterns of long-term recordings into groups of similar patterns. Altered states of brain synchronization were identified as deviating from patterns in the reference library and were evaluated relative to the times of seizure onset in terms of sensitivity and specificity. METHODS: A phase-locking measure was estimated using a sliding window analysis on 15 frequency bands (2Hz steps between 0 and 30Hz), for all pairs of EEG channels in the epileptogenic temporal lobe (14-20 channels), over the entire data sets (total analyzed duration 305h). The preictal identification encompasses three basic stages: (1) a preprocessing stage involving the determination of a reference library of characteristic interictal synchronization patterns using a K-means algorithm, and the identification of discriminant variables differentiating interictal from preictal states, (2) a classification stage of the synchronization pattern via a minimum Mahalanobis distance to the reference patterns, as well as detection of outliers, (3) an evaluation stage of the sensitivity and specificity of the detection by receiver-operating characteristic curves. RESULTS: In most of the cases (36 of 52 seizures, i.e. 70%), a specific state of brain synchronization can be observed several hours before the actual seizure. The changes involved both increases and decreases of the synchronization levels, occurring mostly within the 4-15Hz frequency band, and were often localized near the primary epileptogenic zone. CONCLUSIONS: The analysis of phase synchronization offers a way to distinguish between a preictal state and normal interictal activity. These findings suggest that brain synchronizations are preictally altered in the epileptogenic temporal lobe, inducing a pathological state of higher susceptibility for seizure activity. SIGNIFICANCE: Phase synchronization is capable of extracting information from the EEG that allow the definition of a preictal state. Although the proposed analysis does not constitute genuine seizure anticipation, these changes in neuronal synchronization may provide helpful information for prospective seizure warning.

Brain Mapping↗

Isoflurane disrupts anterio-posterior phase synchronization of flash-induced field potentials in the rat.

Consciousness presumes a set of integrated functions such as sensory processing, attention, and interpretation, and may depend upon both local and long-range phase synchronization of neuronal activity in cerebral cortex. Here we investigated whether volatile anesthetic isoflurane at concentrations that produce loss of consciousness (LOC) disrupts long-range anterio-posterior and local anterior synchronization of neuronal activity in the rat. In six rats, deep electrodes were chronically implanted in the primary visual cortex (V1) and in two areas of the motor cortex (M1 and M2) for recording of intracortical event-related potentials (ERP). Thirty discrete flashes were presented at random interstimulus intervals of 15-45 s, and ERPs were recorded at stepwise increasing isoflurane concentrations of 0-1.1%. Neuronal synchronization was estimated using wavelet coherence computed from the ERP data band-pass filtered at 5-50 Hz. We found that (1) in the waking state, long-range anterio-posterior coherence in 5-25 Hz and 25-50 Hz frequency bands was significantly higher than local anterior coherence; (2) anterio-posterior coherence in both 5-25 Hz and 26-50 Hz bands was significantly reduced by isoflurane in a concentration-dependent manner; (3) local anterior coherence was not affected by isoflurane at any of the concentrations studied. These findings suggest that a disruption of long-range anterio-posterior rather than local anterior synchronization of neuronal activity precedes the anesthetic-induced loss of consciousness.

Anesthetics, Inhalation↗

Intracellular [Cl(-)] modulates synchronous electrical activity in rat neocortical neurons in culture by way of GABAergic inputs.

The influence of GABAergic neurons on spontaneous electrical activities of neocortical neurons in culture, which was estimated to be about 9.5% of the total neurons by immunohistochemistry, was examined using dual whole-cell recording. Synchronized depolarization or hyperpolarization was observed in recorded neurons with pipettes containing low [Cl(-)] solution, while synchronized bursting of action potentials (APs) was observed with pipettes containing high [Cl(-)] solution. Spontaneous currents (SCs) were synchronous in all pairs tested with either pipettes containing low or high [Cl(-)] solution and spontaneous outward currents (SOCs) observed at around -30 mV were sensitive to the GABA-A receptor antagonist, bicuculline. Their reversal potential (V(rev)) was linearly related to the logarithm of Cl(-) activity in the pipette (-56.9 mV/decade). The intracellular chloride concentration was estimated from the V(rev) of SCs with gramicidin perforated-patch recordings and was between 5.9 and 28.1 mM (mean: 13.0 mM). These results suggest that GABA depolarized some neurons and hyperpolarized others, depending on the E(Cl). Bicuculline decreased the frequency of periodic depolarized potentials and increased their amplitudes. However, perfusion with low [Cl(-)] bath solution did not decrease the frequency. Our data indicate that recurrent subthreshold electrical activities by GABAergic inputs along with glutamatergic inputs take part in deterring synchronized bursting and that intracellular [Cl(-)] can modulate this bursting.

Action Potentials↗

Coherent oscillations in membrane potential synchronize impulse bursts in central olfactory neurons of the crayfish.

Lateral protocerebral interneurons (LPIs) in the central olfactory pathway of the freshwater crayfish Procambarus clarkii reside within the lateral protocerebrum and receive direct input from projection neurons of the olfactory midbrain. The LPIs exhibit periodic (0.5 Hz) changes in membrane potential that are imposed on them synaptically. Acute surgical experiments indicate that the synaptic activity originates from a group of oscillatory neurons lying within the lateral protocerebrum. Simultaneous intracellular recordings from many LPI pairs indicate that this periodic synaptic input is synchronous and coherent among the population of approximately 200 LPIs on each side of the brain. In many LPIs, specific odors applied to antennules in isolated head preparations generate long-lasting excitatory postsynaptic potentials and impulse bursts. The impulse bursts are generated only near the peaks of the ongoing depolarizations, approximately 1 s after stimulus application, and so the periodic baseline activity is instrumental in timing burst generation. Simultaneous recordings from pairs of LPIs show that, when impulse bursts occur in both cells after an odorant stimulus, they are synchronized by the common periodic depolarizations. We conclude that the common, periodic activity in LPIs can synchronize impulse bursts in subsets of these neurons, possibly generating powerful long-lasting postsynaptic effects in downstream target neurons.

Animals↗