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Long-term stability and consistency of EEG event-related (de-)synchronization across different cognitive tasks.

OBJECTIVE: We examined whether task-related band power changes (event-related desynchronization/synchronization; ERD/ERS) that have been linked to individual differences in cognitive ability demonstrate satisfying temporal stability and cross-situational consistency. METHODS: Multi-channel EEG recordings from 29 adults, assessed at three different occasions over 2 years were examined. Between-session correlations and consistency coefficients (Cronbach's alpha) across the three experiments were evaluated for both, spectral power features of the resting EEG and ERD/ERS estimates while the participants performed some cognitive task (i.e. different elementary cognitive tasks that put comparable demands on the participants). RESULTS: ERD/ERS values, while subjects performed a cognitive task, demonstrated satisfactory stability and consistency (i.e. >0.7), whereby the degree of consistency varied as a function of frequency band and brain region. Highest consistency was found for the 8-10 Hz ERD in parieto-occipital recording sites (i.e. >0.9). In resting EEG, mean alpha (gravity) frequency was the most stable EEG feature. CONCLUSIONS: The present data suggest that ERD/ERS phenomena in different narrow frequency bands are rather stable over time and across different situations. The relatively high reproducibility of ERD/ERS promotes the usefulness of this measure in assessing individual differences of physiological activation patterns accompanying cognitive performance. SIGNIFICANCE: This study addresses the issue of reproducibility of EEG in general and ERD/ERS experiments in particular, which is a prerequisite for both basic research and clinical studies.

Adolescent↗

Increased neuronal communication accompanying sentence comprehension.

The main purpose of this study was to examine large-scale oscillatory activity and frequency-related neuronal synchronization during the comprehension of English spoken sentences of different complexity. Therefore, EEG coherence during the processing of subject-subject (SS)- and more complex subject-object (SO)-relatives was computed using an adaptive fitting approach of bivariate auto-regressive moving average (ARMA) models which enabled the continuous calculation of coherence in the course of sentence processing with a high frequency resolution according to the dynamic changes of the EEG signals. Coherence differences between sentence types were observed in the theta (4-7 Hz), beta-1 (13-18 Hz) and gamma (30-34 Hz) frequency ranges, though emerging during the processing of different parts of these sentences: gamma differences were evident mainly during the relative clause while theta and beta-1 differed significantly following the end of the relative clause. These findings reveal no simple one to one map between EEG frequencies and cognitive operations necessary for sentence comprehension. Instead, they indicate a complex interplay and dynamic interaction between different EEG frequencies and verbal working memory, episodic memory, attention, morpho-syntactic and semantic-pragmatic analyses, which though distinct often co-occur.

Adult↗

Primary role of the serotonergic midline system in synchronized spontaneous activity during development of the embryonic mouse hindbrain.

In the developing embryonic mouse hindbrain, we have shown that previously widespread synchronized spontaneous activity at E11.5 retracts to the initiating zone of the rostral hindbrain by E13.5, and ceases completely by E14.5. We now confirm that at E11.5 and E13.5, the primary driver of spontaneous activity is serotonergic input, while other transmitters (GABA, glutamate, NE, and ATP) have only modulatory roles. Using immunocytochemistry, we also show that at E13.5, 5-HT-positive neurons in the midline extend over a larger rostro-caudal distance than at E11.5, and that in the presumptive initiating zone, cell bodies occupy a band that extends 200 microm laterally on each side of the midline, with extensive axonal processes. The 5-HT2A receptor retains expression in lateral tissue over this developmental time. We find that in addition to being sensitive to 5-HT receptor antagonists, spontaneous activity is also abolished by blockers of gap junctions, and is increased in frequency and lateral spread by application of ammonium, presumably via increased intracellular pH augmenting gap junction conductance. Thus, 5-HT neurons of the midline remain the primary drivers of spontaneous activity at several stages of development in the hindbrain, relying in part on gap junctional communication during initiation of activity.

Animals↗

Neuromagnetic correlates of sensorimotor synchronization.

Sensorimotor synchronization tasks, in which subjects have to tap their finger in synchrony with an isochronous auditory click, typically reveal a synchronization error with the tap preceding the click by about 20 to 50 msec. Although extensive behavioral studies and a number of different explanatory accounts have located the cause of this so-called "negative asynchrony" on different levels of processing, the underlying mechanisms are still not completely understood. Almost nothing is known about the central processes, in particular, which sensory or motor events are synchronized by subjects. The present study examined central-level processing in synchronization tasks with magnetoencephalography (MEG). Eight subjects synchronized taps with their right index finger to an isochronous binaural pacing signal presented at an interstimulus interval of 800 msec. To gain information on central temporal coupling between "tap" and "click," evoked responses were averaged time-locked to the auditory signal and the tap onset. Tap-related responses could be explained with a three dipole model: One source, peaking at approximately 77 msec before tap onset, was localized in contralateral primary motor cortex (MI); the two other sources, peaking approximately at tap onset and 75 msec after tap onset, in contralateral primary somatosensory cortex (SI). Temporal coupling of these sources was compared in relation to different trigger points. The second SI source was equally well time-locked to the tap and to the auditory click. Furthermore, analysis of the time locking of this source activity as a function of the temporal order of tap and click showed that the second event - irrespective whether tap or click - was decisive in triggering the second SI source. This suggests that subjects use mainly sensory feedback in judging and evaluating whether they are "keeping time."

Acoustic Stimulation↗

Neural encoding of single-formant stimuli in the ventral cochlear nucleus of the chinchilla.

Responses of the principal unit types in the ventral cochlear nucleus of the chinchilla were studied with a single-formant stimulus set that covered fundamental frequency (f0) from 100 Hz to 200 Hz and formant center frequency (F1) from 256 to 782 Hz. Temporal coding for f0 and F1 was explored for 95 stimulus combinations of f0 (n = 5) and F1 (n = 19) in primarylike, onset and chopper unit categories. Several analyses that explored temporal coding were employed including: autocorrelation, interspike interval analysis, and synchronization to each harmonic of f0. In general, the representation of f0 is better in onset and chopper units than in primarylike units. Nearly all units in the cochlear nucleus showed a gain in phase locking to the envelope (f0) of the single-formant stimulus relative to the auditory nerve. The fundamental is represented directly in neural discharges of units in the cochlear nucleus with an interval code (also Cariani and Delgutte, 1996; Rhode, 1995). The formant is represented in the temporal domain in primarylike units, though some chopper and onset units also possess the ability to code F1 through discharge synchrony. Onset-I units, which are associated with the octopus cells, exhibited the strongest phase locking to f0 of any unit types studied. The representation of f0 and F1 in the temporal domain is weak or absent in some units. All-order-interspike interval distributions computed for populations of units show preservation of temporal coding for both f0 and F1. Results are in agreement with earlier amplitude modulation studies that showed nearly all cochlear nucleus unit types phase lock to the signal envelope better than auditory nerve fibers over a considerable range of signal amplitudes.

Acoustic Stimulation↗

Spatial restriction of spontaneous activity towards the rostral primary initiating zone during development of the embryonic mouse hindbrain.

In the developing embryonic mouse hindbrain, we have previously shown that synchronized spontaneous activity is driven by midline serotonergic neurons at E11.5. This is mediated, at least in part, by the 5-HT2A receptor, which is expressed laterally in the hindbrain. Activity at E11.5 is widespread within the hindbrain tissue, propagating from the midline to more lateral regions. Using rapid acquisition of [Ca2+]i events along the midline, we now show that the rostral midline, primarily in the region of former rhombomere r2, is the primary initiating zone for this activity. We propose that at E11.5, the combined events along the rostral-caudal axis in combination with events propagating along the medial-lateral axis could assign positional information to developing neurons within the hindbrain. With further development, to E13.5, both the lateral and caudal dimensions of spontaneous activity retract to the rostral midline, occupying an area only 14% of that exhibited at E11.5. We also show that increased levels of [K+]o (to 8 mM) at E13.5 are able to increase the spread of spontaneous activity laterally and rostro-caudally. This suggests that spontaneous activity in the hindbrain depends in a dynamic way on the dominant initiating zone of the rostral midline, and that this relationship changes over development.

Animals↗

Event-related desynchronization and synchronization during a memory task in children.

OBJECTIVE: To examine the event-related desynchronization (ERD) and synchronization (ERS) responses of several narrow electroencephalographic (EEG) frequency bands in children during an auditory memory task. METHODS: ERD/ERS responses of the 4-6, 6-8, 8-10 and 10-12 Hz EEG frequency bands were studied in 12 children (mean age 12 years) while they performed an auditory memory task. Twelve adult subjects served as a control group. RESULTS: The children's ERD/ERS responses differed from those of the adult's in the 4-6, 6-8 and 8-10 Hz EEG frequency bands, especially during retrieval from memory. The children's 4-6 Hz initial ERS responses were of lesser amplitude and of delayed latency as compared to those of the adults. In the 6-8 and 8-10 Hz frequency bands, especially during retrieval from memory, the children's ERD responses were of lesser magnitude than those of the adults. In the 10-12 Hz frequency band, no differences were observed between the ERD/ERS responses between the children and adults. CONCLUSIONS: Our results suggest that theta and alpha response systems might participate in auditory information processing already at this age, although not being fully developed. Memory systems involving retrieval may be the last to mature.

Acoustic Stimulation↗

Decreased coherence in higher frequency ranges (beta and gamma) between central and frontal EEG in patients with schizophrenia: A preliminary report.

Schizophrenia is associated with a dysfunction of cognitive integration that may be due to abnormalities in inhibitory neural circuitry. A previous study found a failure of gamma band (25-45 Hz) synchronization in patients with schizophrenia compared to controls. Another recent study also stressed the importance of investigating high frequencies in the scalp-recorded sleep electroencephalogram (EEG). In this study, we compared coherence between first episode drug-naïve patients with schizophrenia (n=8) and age- and sex-matched normal controls (n=8) using two 32-s epochs of C4 and F4 EEG. The coherence was obtained using 4096 data points (128 Hz signal) using cross-spectral analysis with Blackman-Tukey window in beta (15.25-24.75 Hz) and gamma (25-44.75 Hz) frequency bands. We used wake, non-rapid eye movement (NREM) and rapid eye movement (REM) sleep periods for the analyses. Our results show a significant decrease in coherence in both beta and gamma frequency bands in patients. Post-hoc 't' tests revealed a significantly lower coherence only during the wake stage in patients with schizophrenia in beta as well as gamma frequency bands. These results further support the importance of the analyses of high-frequency bands in the EEG and support previous findings of abnormal neural synchrony in patients with schizophrenia. These results have been discussed further in relation to wake and sleep stages.

Adult↗

Nucleus basalis magnocellularis and pedunculopontine tegmental nucleus: control of the slow EEG waves in rats.

Certain human disorders, which are characterized by learning and memory disturbances, are also accompanied by synchronizations of the cortical electroencephalogram (EEG). Although the EEG synchronization was casually related to the degeneration of the cholinergic basal forebrain, degenerations were found also in the cholinergic pontomesencephalic tegmentum. The present study was carried out to prove whether lesions of rat's pedunculopontine tegmental nucleus (PPTg) may cause EEG synchronizations like lesions of the basal forebrain. The effects of the unilateral ibotenic acid lesions of the nucleus basalis magnocellularis (NBM) and of the PPTg on the frontal and occipital EEG were compared in different behavioural states. The NBM lesion caused unilateral spectral power increases of all frequencies up to 20 Hz in the frontal EEG that were stronger with increased arousal level. Additionally, synchronized spike and wave discharges appeared in the frontal EEG. The NBM may suppress especially the delta EEG waves in the frontal motor cortex during motor active behaviours. The PPTg lesion caused unilateral suppressions of the occipital theta rhythm during exploratoring sniffing. During grooming the power of the frontal delta waves were elevated. Furthermore, the PPTg lesion caused a suppression of slow sleep waves. The results may indicate that a degeneration of both brain regions may change differently the power of the cortical slow EEG waves.

Acetylcholinesterase↗

Extrinsic modulation of theta field activity in the entorhinal cortex of the anesthetized rat.

Field recordings of the entorhinal cortex (EC) were studied and compared to those recorded concomitantly in the dentate region of the hippocampal formation (HPC) in the urethane anesthetized rat. The EC, like the HPC, showed two main variations of spontaneous field activity: a desynchronized, large amplitude irregular activity and a synchronized, rhythmic, slow frequency field activity (RSA or theta). Corroborating previous research, a phase reversal was seen across layer II of the EC and when recorded superficial to this layer, EC theta was phase-locked to that recorded from the HPC (dentate). Entorhinal cortex (and HPC) theta could be evoked by the application of moderate tail pinches (sensory stimulation), by pharmacological treatments enhancing cholinergic transmission, and by electrical stimulation of the posterior hypothalamus. Spectral analysis revealed that in all cases, theta was produced coherently across the two limbic structures. Entorhinal cortex (and HPC) production of theta could be abolished by pharmacological treatments disrupting cholinergic transmission, and by reversible procaine inactivation of the medial septal region. Therefore, it was concluded that limbic theta is modulated spontaneously, and with sensory and hypothalamic stimulation through the activity of cells in the medial septal region via muscarinic neurotransmission. It was also hypothesized that the activation of cells in the posterior hypothalamus linearly codes the frequency, and to a lesser extent the power, of EC and HPC theta. Given these findings and the coincidence and coherence of the occurrence of theta across the EC and HPC, it was postulated that it occurs via a parallel mechanism in the two areas.

Anesthesia↗

Event-related desynchronization/synchronization during an auditory-verbal working memory task in mild Parkinson's disease.

OBJECTIVE: Event-related desynchronization (ERD) and synchronization (ERS) were studied during an auditory-verbal working memory task in patients suffering from Parkinson's disease (PD). METHODS: PD patients (n=7, mean age 59) at a mild stage of the disease volunteered in the study. A group of healthy subjects (n=10, mean age 61) served as control group. ERD and ERS of the 1-25 Hz EEG frequencies were studied using wavelet transforms during memory encoding and retrieval. RESULTS: Both groups performed equally well on the memory task. Statistically significant differences in ERD/ERS responses were observed in posterior electrodes during encoding of the memory set due to the fact that alpha ( approximately 10-15 Hz) ERS was elicited in the controls, but not in the PD group. In broad frequency bands ( approximately 6-25 Hz) ERD responses were observed in both groups during memory retrieval. A pre-stimulus alpha ERS seen in the control group was absent in the PD group. CONCLUSIONS: The results of this preliminary study indicate that Parkinson's disease might affect brain oscillatory responses in the alpha frequency range in the encoding phase of auditory-verbal working memory. SIGNIFICANCE: The ERD/ERS patterns may reflect neurophysiological alterations in the processes underlying working memory deficits in PD.

Acoustic Stimulation↗

Decreased theta event-related synchronization during working memory activation is associated with progressive mild cognitive impairment.

BACKGROUND: Among the different quantitative electroencephalographic markers, theta activity is known to reflect neural resources involved in memory processes and directed attention. Previous studies suggested that synchronization likelihood analysis in theta-band frequency might be a sensitive method to identify early alterations of neuronal networks in mild cognitive impairment (MCI). METHODS: We report here a longitudinal study of 24 MCI patients with theta event-related synchronization (ERS) analysis during the n-back working memory task and neuropsychological follow-up after 1 year. Statistical analysis included analysis of variance and logistic regression to assess the relationship between cognitive decline and theta ERS. RESULTS: Upon follow-up, 13 MCI patients showed progressive MCI and 11 remained stable. In both groups, the phasic increase in theta amplitude after stimulus presentation did not depend on working memory load and electrode sites. Progressive MCI cases displayed significantly lower theta ERS power over all electrode sites compared to stable MCI cases. Theta ERS was significantly related to the cognitive outcome explaining 15.5% of its variability. In terms of MCI classification, the best combination of sensitivity and specificity was 0.87 and 0.60, respectively, with an area under the corresponding receiver operating characteristic curve reaching 76%. CONCLUSIONS: The present data indicate that a decrease in the early phasic theta ERS power during working memory activation may predict cognitive decline in MCI. This phenomenon is not related to working memory load but may reflect the presence of early deficits in directed attention-related neural circuits in MCI.

Aged↗

Scene segmentation by spike synchronization in reciprocally connected visual areas. II. Global assemblies and synchronization on larger space and time scales.

We present further simulation results of the model of two reciprocally connected visual areas proposed in the first paper [Knoblauch and Palm (2002) Biol Cybern 87:151-167]. One area corresponds to the orientation-selective subsystem of the primary visual cortex, the other is modeled as an associative memory representing stimulus objects according to Hebbian learning. We examine the scene-segmentation capability of our model on larger time and space scales, and relate it to experimental findings. Scene segmentation is achieved by attention switching on a time-scale longer than the gamma range. We find that the time-scale can vary depending on habituation parameters in the range of tens to hundreds of milliseconds. The switching process can be related to findings concerning attention and biased competition, and we reproduce experimental poststimulus time histograms (PSTHs) of single neurons under different stimulus and attentional conditions. In a larger variant the model exhibits traveling waves of activity on both slow and fast time-scales, with properties similar to those found in experiments. An apparent weakness of our standard model is the tendency to produce anti-phase correlations for fast activity from the two areas. Increasing the inter-areal delays in our model produces alternations of in-phase and anti-phase oscillations. The experimentally observed in-phase correlations can most naturally be obtained by the involvement of both fast and slow inter-areal connections; e.g., by two axon populations corresponding to fast-conducting myelinated and slow-conducting unmyelinated axons.

Animals↗

Effects of normal aging on event-related desynchronization/synchronization during a memory task in humans.

Event-related desynchronization (ERD) and event-related synchronization (ERS) of the 1-20Hz EEG frequencies were studied using wavelet transforms in young (n = 10, mean age 22) and elderly subjects (n = 10, mean age 65) performing an auditory Sternberg memory task with words as stimuli. In both age groups, encoding of the four-word memory set elicited ERS in the theta and alpha frequency range. Theta ERS, and ERD in the alpha and beta bands were observed during retrieval. During encoding, the elderly showed greater alpha ERS and smaller theta ERS. During retrieval, smaller alpha ERD and theta ERS was found in the elderly subjects. Also, in the elderly, beta ERD was elicited in the late time window during retrieval. The statistically significant differences between the age groups were more marked during retrieval than during encoding. The results indicate that although the two groups performed equally well behaviorally in the task and the elderly subjects were cognitively intact, normal aging affects oscillatory theta, alpha and beta responses particularly during retrieval from working memory. The ERD/ERS patterns of the elderly resemble those of children found in a recent study, which might suggest that those memory-related brain processes that evolve later in childhood are the first to be affected in older age.

Acoustic Stimulation↗

Synchronization of gamma oscillations increases functional connectivity of human hippocampus and inferior-middle temporal cortex during repetitive visuomotor events.

Do recency processes associated with repetitive sensorimotor events modulate the magnitude and functional coupling of brain rhythmicity in human temporal cortex? Intracranial stereo electroencephalographic activity (SEEG; 256 Hz sampling rate) was recorded from hippocampus, and inferior (BA20) and middle (BA21) temporal cortex in four epilepsy patients. The repetitive events were represented by predicted imperative somatosensory stimuli (CNV paradigm) triggering hand movements ("repetitive visuomotor") or counting ("repetitive counting"). The non-repetitive events were "rare" (P3 paradigm) somatosensory stimuli triggering hand movements ("non-repetitive visuomotor") or counting ("non-repetitive counting"). Brain rhythmicity was indexed by event-related desynchronization/synchronization (ERD/ERS) of SEEG data, whereas the functional coupling was evaluated by spectral SEEG coherence between pairs of the mentioned areas. The frequency bands of interest were theta (4-8 Hz), alpha (8-12 Hz), beta (14-30 Hz), and gamma (32-46 Hz). Compared to the non-repetitive events, the "repetitive visuomotor" events showed a significant beta and gamma ERS in the hippocampus and a significant theta ERD in the inferior temporal cortex. Furthermore, the "repetitive visuomotor" events induced a task-specific significant gamma coherence among the examined areas. These results suggest that recency processes do modulate the magnitude and functional coupling of brain rhythmicity (especially gamma) in the human temporal cortex.

Adult↗

Region-specific modulation of electrically induced synchronous oscillations in the rat hippocampus and cerebral cortex.

Strong tetanization induces synchronous membrane potential oscillations (seizure-like afterdischarge) in mature pyramidal cells of the hippocampal CA1 region. To investigate whether local networks in other brain regions can generate such an afterdischarge independently, we studied the inducibility of afterdischarge in individual 'isolated slices' of the rat hippocampal CA1 and CA3 regions, dentate gyrus (DG), entorhinal cortex (EC), and temporal cortex (TC) using intracellular and extracellular recordings. The strong tetanization constantly induced afterdischarges in the CA1 and CA3 pyramidal cells as well as in the EC and TC superficial principal cells. However, parameters of the afterdischarges, such as the frequency and duration of afterdischarges, varied among the regions. A mixture of N-methyl-D-aspartate (NMDA) and non-NMDA receptor antagonists or a GABA(A) receptor antagonist completely blocked the afterdischarges. Local GABA application during the afterdischarge elicited depolarization, rather than hyperpolarization. Moreover, reversal potentials of the afterdischarge were around -40 mV. In contrast, the tetanization resulted in occasional afterdischarge-like activities in DG slices, which were blocked by the non-NMDA or GABA(A) receptor antagonist. These findings suggest that the afterdischarges mediated through the excitatory GABAergic and glutamatergic transmissions might be common to, but be modulated differently by individual local networks in the hippocampus and cortex.

Animals↗

Determination of language dominance with synthetic aperture magnetometry: comparison with the Wada test.

Cerebral dominance for language function was investigated with synthetic aperture magnetometry (SAM). The results were compared with those of the Wada test. SAM is a spatial filtering technique that enables demonstration of the spatiotemporal distribution of oscillatory changes (synchronization and desynchronization) in magnetoencephalography (MEG) signals elicited by specific brain activation. MEG was conducted during a silent reading task in 20 consecutive preoperative neurosurgical patients who also underwent a Wada test. The spatial distribution of oscillatory changes related to silent reading was shown tomographically with SAM as statistical images. Language dominance was estimated by the laterality index, which scales the lateralization of the beta (13-25 Hz) and low gamma (25-50 Hz) band desynchronizations in the inferior frontal gyrus (IFG) or middle frontal gyrus (MFG). Oscillatory changes were distributed multifocally and bilaterally in the occipital cortex, IFG or MFG, and temporo-parieto-occipital border regions. In 19 patients (95%), language lateralization estimated by the laterality index was congruent with the result of the Wada test. In left-handed patients, SAM analysis clearly differentiated language dominance (left, right, or bilateral), and the findings were confirmed by the Wada test. Lateralization of beta or low gamma band desynchronizations in the IFG or MFG is a good indicator of the side of language dominance. Reliability of MEG imaging with SAM is sufficient to evaluate language dominance preoperatively in neurosurgical patients.

Adolescent↗

Basic mechanisms of central rhythms reactivity to preparation and execution of a voluntary movement: a stereoelectroencephalographic study.

OBJECTIVE: To localize the sources of mu, beta and gamma rhythms and to explore the functional significance of their reactivity. METHODS: We used the method of quantification of event-related desynchronization (ERD) and synchronization (ERS) to analyze the reactivity of intracerebral rhythms recorded in stereoelectroencephalography within the sensorimotor areas during the preparation and the execution of a simple self-paced hand movement. We recorded 3 epileptic subjects who were explored before a surgical treatment. RESULTS: An ERD of mu and beta rhythms has been recorded before the movement onset in the precentral gyrus, spreading then to the postcentral gyrus and to the frontal medial cortex. The frontal lateral cortex was inconstantly involved during the movement. The movement offset was followed by an important and focused beta ERS which was found within the pre- and post-central gyrus and the frontal medial cortex. Within the beta band, we observed several narrower bands with different reactivities and locations. Focused gamma reactivity was also found in the precentral and postcentral gyri. CONCLUSIONS: The reactivities of mu and beta rhythms are different but their locations overlap. Mu ERD is a diffuse phenomenon that reflects the activation of all the sensorimotor areas during a simple movement. Beta band is likely to be composed of different rhythms with different functional significance. The primary motor area seems to contain two distinct areas with different reactivity to the movement preparation and execution.

Brain Mapping↗