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Dynamics of spiking neurons connected by both inhibitory and electrical coupling.

We study the dynamics of a pair of intrinsically oscillating leaky integrate-and-fire neurons (identical and noise-free) connected by combinations of electrical and inhibitory coupling. We use the theory of weakly coupled oscillators to examine how synchronization patterns are influenced by cellular properties (intrinsic frequency and the strength of spikes) and coupling parameters (speed of synapses and coupling strengths). We find that, when inhibitory synapses are fast and the electrotonic effect of the suprathreshold portion of the spike is large, increasing the strength of weak electrical coupling promotes synchrony. Conversely, when inhibitory synapses are slow and the electrotonic effect of the suprathreshold portion of the spike is small, increasing the strength of weak electrical coupling promotes antisynchrony (see Fig. 10). Furthermore, our results indicate that, given a fixed total coupling strength, either electrical coupling alone or inhibition alone is better at enhancing neural synchrony than a combination of electrical and inhibitory coupling. We also show that these results extend to moderate coupling strengths.

Action Potentials↗

Exploration of event-induced EEG phase synchronization patterns in cognitive tasks using a time-frequency-topography visualization system.

In this paper, we present a method for the study of synchronization patterns measured from EEG scalp potentials in psychophysiological experiments. This method is based on various techniques: a time-frequency decomposition using sinusoidal filters which improve phase accuracy for low frequencies, a Bayesian approach for the estimation of significant synchrony changes, and a time-frequency-topography visualization technique which allows for easy exploration and provides detailed insights of a particular experiment. Particularly, we focus on in-phase synchrony using an instantaneous phase-lock measure. We also discuss some of the most common methods in the literature, focusing on their relevance to long-range synchrony analysis; this discussion includes a comparison among various synchrony measures. Finally, we present the analysis of a figure categorization experiment to illustrate our method.

Bayes Theorem↗

Heterogeneity among hippocampal pyramidal neurons revealed by their relation to theta-band oscillation and synchrony.

Intracellular recordings were made in the dorsal hippocampal formation of urethane-anesthetized rats as the local field activity spontaneously cycled between a synchronous condition termed theta and an asynchronous condition termed LIA. All cells reported in this study were labeled with Neurobiotin and classified as theta-related or non-theta-related according to the system of Colom and Bland [Colom, L.V., Bland, B.H., 1987. State-dependent spike train dynamics of hippocampal formation neurons: evidence for theta-ON and theta-OFF cells. Brain Res. 422; 277-286]. The findings are the first demonstration that hippocampal pyramidal cells are functionally heterogeneous in relation to the generation of theta-band oscillation and synchrony. In field CA1 pyramidal cells formed theta-related subsets of phasic theta-ON cells and tonic theta-ON cells and non-theta-related subsets of simple spike discharging cells, complex spike discharging cells and "silent" cells. Similar findings were evident for CA3 pyramidal cells.

Animals↗

Sex differences in functional connectivity in first-episode and chronic schizophrenia patients.

OBJECTIVE: There has been consistent evidence for a lower incidence and milder course of schizophrenia in women, yet there have been very few investigations of sex differences in brain function in this disorder. This study used a new high-temporal-resolution measure of functional brain connectivity to test the prediction that female patients would show relatively greater inter- and intrahemispheric connectivity than male patients, particularly in the early stage of schizophrenia. METHOD: Forty patients with chronic schizophrenia (20 women and 20 men) and 24 patients with first-episode schizophrenia (12 women and 12 men) and their respective matched comparison groups completed a conventional auditory oddball task. Phase synchronous gamma (40 Hz) activity was extracted from EEG recording during the task and time-locked to the oddball (target) stimuli. RESULTS: Chronic schizophrenia subjects showed a reduction in global functional connectivity (lower gamma phase synchrony) relative to their matched healthy subjects. Unexpectedly, this reduction was most apparent in female patients. By contrast, while first-episode patients showed a general reduction in the speed of frontal connectivity, the speed of global connectivity was relatively faster in female patients. CONCLUSIONS: This is the first study to investigate sex differences in schizophrenia that used the functional connectivity measure of gamma phase synchrony. The results suggest that in female patients with schizophrenia, additional breakdowns in brain network connectivity may develop with illness chronicity.

Adult↗

Oscillatory synchronization requires precise and balanced feedback inhibition in a model of the insect antennal lobe.

In the insect olfactory system, odor-evoked transient synchronization of antennal lobe (AL) projection neurons (PNs) is phase-locked to the oscillations of the local field potential. Sensory information is contained in the spatiotemporal synchronization pattern formed by the identities of the phase-locked PNs. This article investigates the role of feedback inhibition from the local neurons (LNs) in this coding. First, experimental biological results are reproduced with a reduced computational spiking neural network model of the AL. Second, the low complexity of the model leads to a mathematical analysis from which a lower bound on the phase-locking probability is derived. Parameters involved in the bound indicate that PN phase locking depends not only on the number of LN-evoked inhibitory postsynaptic potentials (IPSPs) previously received, but also on their temporal jitter. If the inhibition received by a PN at the current oscillatory cycle is both perfectly balanced (i.e., equal to the mean inhibitory drive) and precise (without any jitter), then the PN will be phase-locked at the next oscillatory cycle with probability one.

Animals↗

Electrophysiological, behavioral, and subjective indexes of workload when performing multiple tasks: manipulations of task difficulty and training.

This study examined whether alpha event-related desynchronization (ERD) and theta event-related synchronization (ERS) could successfully measure changes in cognitive workload and training while an operator was engaged in a continuous, interactive, control task(s). Alpha 1 (8-10 Hz) ERD, alpha 2 (10-12 Hz) ERD, and theta (3-7 Hz) ERS were determined for a communications event that occurred during multiple task workload conditions or as a single task. Other measures (alpha and theta EEG power, heart rate, respiration, eye blinks, behavioral performance, and subjective workload ratings) were also evaluated. Results showed that alpha 2 EEG, heart rate, behavioral, and subjective measures were sensitive to changes in workload in the multiple tasks. In addition, eye blink rate and behavioral measures were sensitive to training. Alpha ERD and theta ERS were not sensitive to workload and training in our interactive, multiple task environment. However, they were effective indexes of cognitive/behavioral demands within an interactive single task.

Adult↗

Spectral dynamics of electroencephalographic activity during auditory information processing.

Dynamics of electroencephalographic (EEG) activity during auditory information processing were evaluated in response to changes in stimulus complexity, stimulus discriminability and attention using the oddball paradigm. In comparison to pre-stimulus baseline conditions, auditory stimulation synchronized EEG activity in delta, theta and alpha frequency bands. Event-related synchronization (ERS) effects were greatest at approximately 3 Hz (theta frequency band), and their magnitude depended on stimulus and task demands. Event-related desynchronization (ERD) of EEG activity was observed in the beta frequency band. This effect was greatest at approximately 21 Hz but occurred only for easily discriminable stimuli in attention-related target conditions. Because active discrimination tasks also required a button-press response with the right hand, ERDs involved more complex responses that may be related to a combination of perceptual, motor and cognitive processes. These results demonstrate that oddball and attention-related EEG responses to auditory stimulation could be characterized in the frequency domain. The specific design and analysis features described herein may prove useful since they provide a simple index of the brain's response to stimulation while at the same time provide powerful information not contained in typical time domain analysis.

Adult↗

Does neuronal synchrony underlie visual feature grouping?

Previous research suggests that synchronous neural activity underlies perceptual grouping of visual image features. The generality of this mechanism is unclear, however, as previous studies have focused on pairs of neurons with overlapping or collinear receptive fields. By sampling more broadly and employing stimuli that contain partially occluded objects, we have conducted a more incisive test of the binding by synchrony hypothesis in area MT. We find that synchrony in spiking activity shows little dependence on feature grouping, whereas gamma band synchrony in field potentials can be significantly stronger when features are grouped. However, these changes in gamma band synchrony are small relative to the variability of synchrony across recording sites and do not provide a robust population signal for feature grouping. Moreover, these effects are reduced when stimulus differences nearby the receptive fields are eliminated using partial occlusion. Our findings suggest that synchrony does not constitute a general mechanism of visual feature binding.

Animals↗

Epilepsy with electrical status epilepticus during slow sleep and secondary bilateral synchrony.

In 3 children with "epilepsy with electrical status epilepticus during slow sleep" (ESES), we estimated interhemispheric small time differences (TDs) during spike-wave activity in EEG by coherence and phase analysis by the two-dimensional autoregressive model to differentiate their continuous diffuse spike-waves during slow-wave sleep (CSWS) between primary bilateral synchrony and secondary bilateral synchrony (SBS). Maximal TDs at onset of apparently bilateral synchronous spike-wave bursts (BSSWs) during slow-wave sleep were 12.0-26.5 ms (mean 20.3 ms) with consistent leading hemispheres in eight bursts of the 3 patients, indicating SBS as pathophysiology of their CSWS. This suggestion was supported by their clinico-EEG findings, including the effect of a single oral dose of clobazam (CLB) on EEG. Three ictal BSSWs of atypical absence seizures in 2 patients were also analyzed to obtain maximal TDs of 17.9-41.7 ms (mean 26.3 ms) at onset, with the same leading sides as in sleep, also indicating SBS. Examination of intraburst TD variations showed no consistent disappearance of TDs during the latter part of the bursts, in either sleep or the ictal EEGs of atypical absences, and a role of the corpus callosum was suggested in the generation of SBS in ESES.

Age Factors↗

The role of hippocampal commissures in the interhemispheric transfer of epileptiform afterdischarges in the rat: a study using linear and non-linear regression analysis.

The role of the forebrain commissures and the septal area in the interhemispheric transfer of hippocampal afterdischarges (ADs) was investigated in the rat under halothane anesthesia. Electrical seizures were elicited from the dorsal hippocampus before and after commissurotomy. The degree of relatedness between EEG signals recorded from homologous sites of both hippocampi was quantified using two approaches: (i) a time domain analysis considering an AD as a succession of discrete bursts; the onset times of such bursts were measured and used to estimate interhemispheric onset delays; (ii) using signal analysis the linear (r2) and non-linear (h2) regression coefficients between pairs of EEG signals were computed as a function of time shift between the two signals. In this way the values of association (linear and non-linear) and the corresponding time delays were measured. In general a tetanus applied unilaterally to the dorsal CA3 field resulted in bilaterally synchronous ADs. The estimated interhemispheric time delay was in most cases zero. This bilateral synchrony disappeared after section of a specific part of the ventral hippocampal commissure (VHC), the dorso-caudal third, but was not affected by section of other commissural fibers or by a lesion of the septal area. This study also allowed evaluation of different methods of quantification of the association between EEG signals, namely the linear (r2) and the non-linear (h2) regression coefficients. The latter was shown to be a more robust measure than the former and to yield values of association even in cases in which r2 was at noise level. The experimental findings allow the conclusion that ADs elicited from an epileptogenic focus spread to homologous sites in the contralateral hemisphere following commissural systems that may be strong enough to ensure the forming of one bilateral oscillating system.

Animals↗

Pattern separation and synchronization in spiking associative memories and visual areas.

Scene analysis in the mammalian visual system, conceived as a distributed and parallel process, faces the so-called binding problem. As a possible solution, the temporal correlation hypothesis has been suggested and implemented in phase-coding models. We propose an alternative model that reproduces experimental findings of synchronized and desynchronized fast oscillations more closely. This model is based on technical considerations concerning improved pattern separation in associative memories on the one hand, and on known properties of the visual cortex on the other. It consists of two reciprocally connected areas, one corresponding to a peripheral visual area (P), the other a central association area (C). P implements the orientation-selective subsystem of the primary visual cortex, while C was modeled as an associative memory with connections formed by Hebbian learning of all assemblies corresponding to stimulus objects. Spiking neurons including habituation and correlated noise were incorporated as well as realistic synaptic delays. Three learned stimuli were presented simultaneously and correlation analysis was performed on spike recordings. Generally, we found two states of activity: (i) relatively slow and unordered oscillations at about 20-25 Hz, synchronized only within small regions; and (ii) faster and more precise oscillations around 50-60 Hz, synchronized over the whole simulated area. The neuron groups representing one stimulus tended to be simultaneously in either the slow or the fast state. At each particular time, only one assembly was found to be in the fast state. Activation of the three assemblies switched on a time scale of 100 ms. This can be interpreted as self-generated attention switching. On the time scale corresponding to gamma oscillations, cross correlations between local neuron groups were either modulated or flat. Modulated correlograms resulted if the groups coded features corresponding to a common object. Otherwise, the correlograms remained flat. This behavior is in agreement with experimental results, while phase-code models would generally predict modulated correlations also in the case of different objects. Furthermore, we derive a technical version from our biological associative memory model that accomplishes fast pattern separation parallel in O(log2 n) steps for n neurons and sparse coding.

Action Potentials↗

Brain oscillatory 1-30 Hz EEG ERD/ERS responses during the different stages of an auditory memory search task.

Event-related desynchronization (ERD) and event-related synchronization (ERS) responses of 1-30 Hz EEG frequencies during the different stages of an auditory Sternberg memory task were examined. The ERD/ERS responses were examined separately for successive memory set items (four) and for the two recognition conditions (YES/NO). The presentation of the memory set elicited ERS responses in the theta and alpha frequencies, and also beta ERD responses. These ERD/ERS responses elicited during encoding were found to evolve with successive memory set item presentation. The ERD/ERS responses elicited during the presentation of the probe dissociated significantly between the two recognition conditions (YES/NO). When the probe was included in the memory set (YES condition), recognition elicited stronger alpha and beta frequency ERD responses as compared to the NO condition. The findings from the current study verify that alpha ERD/ERS responses robustly dissociate between auditory encoding and recognition. The increasing alpha ERS responses with increasing memory set item presentation during encoding may be correlates of the functioning phonological loop, active memory maintenance and/or attention. The alpha ERD responses during recognition are undoubtedly associated with auditory memory search processes and distinguish between previously presented versus not presented verbal material. We propose that alpha ERD/ERS responses reflect explicitly auditory memory processes, discriminating between auditory encoding and recognition. Theta ERS responses may be associated with working memory processes, and possibly more specifically with the functioning of the central executive. Beta ERD/ERS responses may reflect also cognitive and/or memory processing, rather than merely the activity of the motor cortices.

Acoustic Stimulation↗

Brain electrical activity in patients with presenile and senile dementia of the Alzheimer type.

Neurophysiological and behavioral data obtained from 9 patients with presenile dementia and 10 with senile dementia of the Alzheimer type were compared with similar data from 25 age- and sex-equivalent controls. Compared with the healthy controls, both patients groups demonstrated increased background electroencephalographic slowing with a reduction in fast activity (synchronization). Topographic analyses of data from electroencephalographic and evoked potential studies indicate that areas of maximal group differences between the presenile patients and their controls include the right posterior temporal and, to a lesser extent, left midtemporal to anterior temporal areas, whereas the maximal differences between the senile patient group and their controls involve the midfrontal and anterior frontal lobes, bilaterally. Moreover, right-sided numerical features derived from topographic maps proved most useful in differentiating the presenile patients and their age-matched controls, whereas bilateral features were more useful in separating senile patients from their controls. These topographic dissimilarities between patient groups suggest that an age-disease interaction exists between patients with presenile and senile dementia of the Alzheimer type. Correlational analyses between neuropsychological test scores and neurophysiological features indicate that increased slowing and decreased fast activity were associated with poorer test performance.

Adult↗

The effect of the Relaxodont brain wave synchronizer on endodontic anxiety: evaluation by galvanic skin resistance, pulse rate, physical reactions, and questionnaire responses.

The effects of a brain wave synchronizer (BWS) on endodontic (root canal) anxiety was evaluated in the clinical practices of the senior author. The experimental groups were: (1) a verbal method (routine calming words by dentist) plus BWS (N = 10) and (2) verbal method plus BWS and alpha relaxation tape (N = 10). The control group was verbal method alone (N = 10). All three groups were evaluated during a complete endodontic treatment by the use of galvanic skin resistance (GSR), pulse rate (PR), physical responses, and pre- and post-treatment questionnaires. Recordings were made during the following periods: (1) start; (2) local anesthetic injection; (3) rubber dam application; (4) drilling; (5) x-ray taking; (6) instrumentation; (7) obturation; and (8) conclusion. Results showed that the experimental groups were significantly better than the control group in the reduction of endodontic anxiety. The findings reinforced the belief that local anesthetic injection is the most anxiety producing aspect of endodontic treatment.

Adult↗

Small-world networks and functional connectivity in Alzheimer's disease.

We investigated whether functional brain networks are abnormally organized in Alzheimer's disease (AD). To this end, graph theoretical analysis was applied to matrices of functional connectivity of beta band-filtered electroencephalography (EEG) channels, in 15 Alzheimer patients and 13 control subjects. Correlations between all pairwise combinations of EEG channels were determined with the synchronization likelihood. The resulting synchronization matrices were converted to graphs by applying a threshold, and cluster coefficients and path lengths were computed as a function of threshold or as a function of degree K. For a wide range of thresholds, the characteristic path length L was significantly longer in the Alzheimer patients, whereas the cluster coefficient C showed no significant changes. This pattern was still present when L and C were computed as a function of K. A longer path length with a relatively preserved cluster coefficient suggests a loss of complexity and a less optimal organization. The present study provides further support for the presence of "small-world" features in functional brain networks and demonstrates that AD is characterized by a loss of small-world network characteristics. Graph theoretical analysis may be a useful approach to study the complexity of patterns of interrelations between EEG channels.

Aged↗

Statistical significance of coincident spikes: count-based versus rate-based statistics.

Inspired by different conceptualizations of temporal neural coding schemes, there has been recent interest in the search for signs of precisely synchronized neural activity in the cortex. One method developed for this task is unitary-event analysis. This method tests multiple single neuron recordings for short epochs with significantly more coincident spikes than expected from independent neurons. We reformulated the statistical test underlying this method using a coincidence count distribution based on empirical spike counts rather than on estimated spike probabilities. In the case of two neurons, the requirement of stationary firing rates, originally imposed on both neurons, can be relaxed; only the rate of one neuron needs to be stationary, while the other may follow an arbitrary time course. By analytical calculations of the test power curves of the original and the revised method, we demonstrate that the test power can be increased by a factor of two or more in physiologically realistic regimes. In addition, we analyze the effective significance levels of both methods for neural firing rates ranging between 0.2 Hz and 30 Hz.

Action Potentials↗

Independent component analysis compared to Laplacian filtering as "deblurring" techniques for event related desynchronization/synchronization.

OBJECTIVES: The aim of the work was to compare two different approaches - one model-dependent, the other data-dependent - for "deblurring" EEG data, in order to improve the estimation of Event-Related Desynchronization/Synchronization. METHODS: Realistic Surface Laplacian filtering (SL) and Infomax Independent Component Analysis (ICA) were applied on multivariate scalp EEG signals (SL: 128 electrodes with MRI-based realistic modeling; ICA: a subset of 19 electrodes, no MRI) prior to beta Event Related Synchronization (ERS) estimation after finger movement in 8 normal subjects. ERS estimation was performed using standard band-pass filtering. ERS peak amplitudes and latencies in the most responsive channel were calculated and the effect of the two methods above was evaluated by one-way analysis of variance (ANOVA) and Sheffe's test. RESULTS: Both methods and their combination significantly improved ERS estimation (greater ERS peak amplitude, p <0.05). The results obtained after ICA on 19 electrodes were not significantly different than the ones obtained with Realistic SL using 128 electrodes and MRI for scalp modeling (p >0.89). CONCLUSIONS: The "low cost" of ICA (19 electrodes, no MRI) imposes such method as a valid alternative to SL filtering. The employ of ICA after SL filtering suggests that the "ideal EEG deblurring method" would unify the two approaches, depending on both the scalp model and the data.

Artifacts↗

Hippocampal electrical activity and different types of conditioned reflexes in dogs.

Changes in hippocampal activity and heart rate were studied in dogs during elaboration of defensive and alimentary conditioned reflexes and switching-over between different types of reinforcement. Conditional stimuli enhanced synchronization and increased frequency of theta wave trains in hippocampal electrical activity. Parallelly to emotional tension of the dogs a significant rise in amplitude of integrated theta rhythm appeared. An increase in the positive correlation between the changes in amplitude of integrated theta rhythm and in heart rate shows that the hippocampal theta rhythm may be used as a correlate of emotional tension. Changes in hippocampal electrical activity in the early stages of formation of conditioned connections and throughout the experiment with conditioned "switching-over" suggest that emotional tension arises under conditions of pragmatic uncertainty when the animal does not have sufficient information for the organization of adaptive reactions. Mom marked changes in heart rate and in amplitude of hippocampal theta rhythm during the elaboration of defensive CRs compared with those observed during the development of response eliminating nociceptive stimulation of partner, suggest that the magnitude of emotional tension depends on the conditioned excitation underlying the response being elaborated.

Animals↗