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Theta waves and behavioral manifestations of alertness and dreaming activity in the rat.

Electro-oscillographic recordings show that when rats wake up, behavioral manifestations such as head dorsal extension and snout and eye movements (which occur in this sequence), occasionally followed by brief ear and limb twitches, are preceded by and are simultaneous with theta waves (around 6.5 Hz in frequency) in neocortical areas 3 and 17 and in the hippocampus. Heart rate increases when the animal passes from synchronized sleep to relaxed wakefulness to exploratory behavior. During desynchronized sleep theta waves, similar to those found during alertness, usually precede and are simultaneous with rostrum and eye movements and brief ear and limb twitches. Heart rate also increases during dreaming activity. A high degree of coherence (near 1.0) was found among theta bursts in different hippocampal areas in both highly attentive alertness and dreaming. It is suggested that theta waves are related to attention as part of some kinds of behavior and of the process of dream shaping and expression, both characterized by consciousness.

Animals↗

[Rhythmic activity of central portions of the cat visual analyzer during different forms of behavior].

Synchronized EEG activity of visual cortex and some other parts of the brain was recorded in cats during milk licking. The frequency range, cortico-subcortical topography and some conditions of its appearance and blockade were analyzed. It was found, that this activity consisted of two rhythms with frequencies of 6--9 and 9--15 c/s and was strongly influenced by visual conditions. By come parameters it also differed from EEG of drowsiness and light sleep. Possible mechanisms and localization of pacemakers of this activity was discussed as well as its similarity to human alpha rhythm.

Animals↗

Automated frequency analysis of synchronous and diffuse sleep spindles.

BACKGROUND: Sleep spindles have different properties in different localizations in the cortex. OBJECTIVES: First main objective was to develop an amplitude-independent multi-channel spindle detection method. Secondly the method was applied to study the anteroposterior frequency differences of pure synchronous (visible bilaterally, either frontopolarly or centrally) and diffuse (visible bilaterally both frontopolarly and centrally) sleep spindles. METHODS: A previously presented spindle detector based on the fuzzy reasoning principle and a level detector were combined to form a multi-channel spindle detector. RESULTS: The spindle detector had a 76.17% true positive rate and 0.93% false-positive rate. Pure central spindles were faster and pure frontal spindles were slower than diffuse spindles measured simultaneously from both locations. CONCLUSIONS: The study of frequency relations of spindles might give new information about thalamocortical sleep spindle generating mechanisms.

Adult↗

Delayed beta synchronization after movement of the more affected hand in essential tremor.

To investigate the pathomechanism of parkinsonian tremor (PT) and essential tremor (ET) by studying the correlation between tremor asymmetry and post-movement beta synchronization (PMBS) of the human EEG. We recorded the EEG of 10 patients with ET, 10 patients with Parkinson's disease and 10 controls. Subjects pressed an on-off switch in a self-paced manner with the thumb of their less (T+) and more (T++) tremulous hand. After digitalization of the EEG from the Cz, C3, C4 electrodes the movement reactive beta frequency, its maximum peak power value and its latency triggered to movement offset were determined. In ET tremor intensity did not influence the power of PMBS, however it was significantly delayed after the movement of the more tremulous hand. In Parkinson's disease after the movement of the more tremulous hand PMBS power was decreased, but it was not delayed. In controls the side of movement had no effect on the power and latency of the PMBS. The neuronal mechanisms underlying PMBS generation are differently affected in essential tremor and Parkinson's disease. The increase of PMBS latency after movement of the more affected hand in ET indicates possible cortical mechanisms in essential tremor generation.

Adult↗

The role of neural synchronization in the emergence of cognition across the wake-sleep cycle.

Searching for the neural code underlying consciousness and cognition is one of the most important activities in contemporary neuroscience. Research with neuronal oscillations at the level of single-neuron, local cell assemblies, and network system have provided invaluable insights into different mechanisms of synaptic interactions involved in the emergence of cognitive acts. A cognitive neuroscience of conscious experience is gradually emerging from behavioral and neuroimaging studies, which can be successfully complemented with the quantitative EEG findings discussed here. This review is an attempt to highlight the value of state-dependent changes in human neurophysiology for a better understanding of the neurobiological substrate underlying those aspects of cognition drastically affected by sleep states. Recent advances related to synchronization mechanisms potentially involved in brain integration processes are discussed, emphasizing the value of scalp and intracranial EEG recordings at determining local and large-scale dynamics in the human brain. Evidence supporting the critical role of state-dependent synchrony in brain integration comes mainly from studies on the theta and gamma oscillations across the wake-sleep continuum, as revealed by human intracranial recordings. This review blends results from different levels of analysis with the firm conviction that state-dependent brain dynamics at different levels of neural integration can provide a deeper understanding of neurobiological correlates of consciousness and sleep functions.

Animals↗

Dynamics of sparsely connected networks of excitatory and inhibitory spiking neurons.

The dynamics of networks of sparsely connected excitatory and inhibitory integrate-and-fire neurons are studied analytically. The analysis reveals a rich repertoire of states, including synchronous states in which neurons fire regularly; asynchronous states with stationary global activity and very irregular individual cell activity; and states in which the global activity oscillates but individual cells fire irregularly, typically at rates lower than the global oscillation frequency. The network can switch between these states, provided the external frequency, or the balance between excitation and inhibition, is varied. Two types of network oscillations are observed. In the fast oscillatory state, the network frequency is almost fully controlled by the synaptic time scale. In the slow oscillatory state, the network frequency depends mostly on the membrane time constant. Finite size effects in the asynchronous state are also discussed.

Action Potentials↗

Modulation of beta oscillations in the subthalamic area during motor imagery in Parkinson's disease.

Activation of the basal ganglia has been shown during the preparation and execution of movement. However, the extent to which the activation during movement is related to efferent processes or feedback-related motor control remains unclear. We used motor imagery (MI), which eliminates peripheral feedback, to further investigate the role of the subthalamic area in the feedforward organization of movement. We recorded local field potential (LPF) activity from the region of the subthalamic nucleus (STN) in eight patients with Parkinson's disease off dopaminergic medication during performance of a warned reaction time task. Patients were instructed to either extend the wrist [motor execution (ME)], to imagine performing the same task without any overt movement (MI), or, in a subgroup, to perform a non-motor visual imagery (VI) task. MI led to event-related desynchronization (ERD) of oscillatory beta activity in the region of the STN in all patients that was similar in frequency, time course and degree to the ERD occurring during ME. The degree of ERD during MI correlated with the ERD in trials of ME and, like ME, was accompanied by a decrease in cortico-STN coherence, so that STN LFP activity during MI was similar to that in ME. The ERD in ME and MI were both significantly larger than the ERD in VI. In contrast, event-related synchronization (ERS) was significantly smaller in trials of MI, and even smaller in trials of VI, than during ME. The data suggest that the activity in the region of the human STN indexed by the ERD during movement is related to the feedforward organization of movement and is relatively independent of peripheral feedback. In contrast, sensorimotor feedback is an important factor in the ERS occurring in the STN area after completion of movement, consistent with a role for this region in trial-to-trial motor learning or the re-establishment of postural set following movements.

Cortical Synchronization↗

Separating stimulus-locked and unlocked components of the auditory event-related potential.

A new method is evaluated for separating stimulus-locked and unlocked components of auditory event-related EEG activity. The new method uses a regression based subtraction procedure as a way to account for latency and amplitude variability within individual trials. It was applied using the oddball paradigm under conditions of active and passive listening and analyzed as spectral correlations (normalized differences) between post-stimulus epochs of standard, target or deviant stimulus conditions and silent pre-stimulus baseline activity. The regression-subtraction procedure accounted for a greater amount of variance than a method that uses linear subtraction alone. The major component of the response to auditory stimulation was an event-related synchronization in the delta and theta (2-4 Hz) frequency range. Event-related desynchronizations were also observed in the 10 Hz (alpha/mu) and in the 20-30 Hz (beta) frequency range. The regression based subtraction procedure provides better separation of stimulus-locked and unlocked components of event-related EEG activity then linear subtraction alone. Stimulus-locked and unlocked components show different patterns and topographies of effects related to attention and active discrimination. Studying both stimulus-locked and unlocked components of event-related EEG reactivity in the frequency domain provides a more comprehensive account of dynamic brain activity subserving auditory information processing.

Acoustic Stimulation↗

Transient synchrony of distant brain areas and perceptual switching in ambiguous figures.

We studied the relationship between perceptual switching in the Necker cube and long-distance transient phase synchronization in EEG. Transient periods of response related synchrony between parietal and frontal areas were observed. They start 800-600, ms prior to the switch response and occur in pairs. Four types of pairs could be distinguished, two of which are accompanied by transient alpha band activity in the occipital area. The results indicate that perceptual switching processes involve parietal and frontal areas; these are the ones that are normally associated with various cognitive processes. Sensory information in the visual areas is involved in some, but not in all, of switching processes. The intrinsic variability, as well as the participating areas, points to the role of strategic cognitive processes in perceptual switching.

Adult↗

Adaptation to tempo changes in sensorimotor synchronization: effects of intention, attention, and awareness.

Adaptation to tempo changes in sensorimotor synchronization is hypothesized to rest on two processes, one (phase correction) being largely automatic and the other (period correction) requiring conscious awareness and attention. In this study, participants tapped their finger in synchrony with auditory sequences containing a tempo change and continued tapping after sequence termination. Their intention to adapt or not to adapt to the tempo change was manipulated through instructions, their attentional resources were varied by introducing a concurrent secondary task (mental arithmetic), and their awareness of the tempo changes was assessed through perceptual judgements. As predicted, period correction was found to be strongly dependent on all three variables, whereas phase correction depended only on intention.

Adaptation, Physiological↗

Frequency flows and the time-frequency dynamics of multivariate phase synchronization in brain signals.

The quantification of phase synchrony between brain signals is of crucial importance for the study of large-scale interactions in the brain. Current methods are based on the estimation of the stability of the phase difference between pairs of signals over a time window, within successive frequency bands. This paper introduces a new approach to study the dynamics of brain synchronies, Frequency Flows Analysis (FFA). It allows direct tracking and characterization of the nonstationary time-frequency dynamics of phase synchrony among groups of signals. It is based on the use of the one-to-one relationship between frequency locking and phase synchrony, which applies when the concept of phase synchrony is not taken in an extended 'statistical' sense of a bias in the distribution of phase differences, but in the sense of a continuous phase difference conservation during a short period of time. In such a case, phase synchrony implies identical instantaneous frequencies among synchronized signals, with possible time varying frequencies of synchronization. In this framework, synchronous groups of signals or neural assemblies can be identified as belonging to common frequency flows, and the problem of studying synchronization becomes the problem of tracking frequency flows. We use the ridges of the analytic wavelet transforms of the signals of interest in order to estimate maps of instantaneous frequencies and reveal sustained periods of common instantaneous frequency among groups of signal. FFA is shown to track complex dynamics of synchrony in coupled oscillator models, reveal the time-frequency and spatial dynamics of synchrony convergence and divergence in epileptic seizures, and in MEG data the large-scale ongoing dynamics of synchrony correlated with conscious perception during binocular rivalry.

Adult↗

Global and local synchrony of coupled neurons in small-world networks.

Synchronous firing of neurons is thought to play important functional roles such as feature binding and switching of cognitive states. Although synchronization has mainly been investigated so far using model neurons with simple connection topology, real neural networks have more complex structures. Here we examine the behavior of pulse-coupled leaky integrate-and-fire neurons with various network structures. We first show that the dispersion of the number of connections for neurons influences dynamical behavior even if other major topological statistics are kept fixed. The rewiring probability parameter representing the randomness of networks bridges two spatially opposite frameworks: precise local synchrony and rough global synchrony. Finally, cooperation of the global connections and the local clustering property, which is prominent in small-world networks, forces synchrony of distant neuronal groups receiving coherent inputs.

Action Potentials↗

On the nonlearnability of a single spiking neuron.

We study the computational complexity of training a single spiking neuron N with binary coded inputs and output that, in addition to adaptive weights and a threshold, has adjustable synaptic delays. A synchronization technique is introduced so that the results concerning the nonlearnability of spiking neurons with binary delays are generalized to arbitrary real-valued delays. In particular, the consistency problem for N with programmable weights, a threshold, and delays, and its approximation version are proven to be NP-complete. It follows that the spiking neurons with arbitrary synaptic delays are not properly PAC learnable and do not allow robust learning unless RP = NP. In addition, the representation problem for N, a question whether an n-variable Boolean function given in DNF (or as a disjunction of O(n) threshold gates) can be computed by a spiking neuron, is shown to be coNP-hard.

Action Potentials↗

A model of the effects of applied electric fields on neuronal synchronization.

We examine the effects of applied electric fields on neuronal synchronization. Two-compartment model neurons were synaptically coupled and embedded within a resistive array, thus allowing the neurons to interact both chemically and electrically. In addition, an external electric field was imposed on the array. The effects of this field were found to be nontrivial, giving rise to domains of synchrony and asynchrony as a function of the heterogeneity among the neurons. A simple phase oscillator reduction was successful in qualitatively reproducing these domains. The findings form several readily testable experimental predictions, and the model can be extended to a larger scale in which the effects of electric fields on seizure activity may be simulated.

Action Potentials↗

Asymmetry and symmetry in brain waves from dolphin left and right hemispheres: some observations after anesthesia, during quiescent hanging behavior, and during visual obstruction.

Studies of sleep in cetaceans (whales, dolphins, and porpoises), substantiated by electrophysiological data, are rare with the great majority of observations having been made by one group from Russia. This group employed hard-wired recording with low-noise cables for their EEG observations, whereas our report describes behavioral and EEG observations of dolphin sleep using telemetry. Marked asymmetry of the EEG was observed during behavioral sleep posture. At different times synchronized slow waves appeared in both left and right brain hemispheres concurrently with lower voltage, faster, desynchronized EEG activity in the opposite hemisphere. On the other hand, during one brief period of sleep behavior, sleep-like EEG activity appeared on leads from both hemispheres. When the animal was exposed to a loud sound, it woke with lower voltage, faster, relatively symmetrical, desynchronized EEG activity appearing from both hemispheres. Additionally, the EEG appeared relatively desynchronized and symmetrical between the two hemispheres when the animal was awake during recovery from pentothal-halothane anesthesia as well as during waking periods when one or both of the animal's eyes were covered by an opaque rubber suction cup.

Animals↗

Trends in electroencephalographic synchronization across nonrapid eye movement sleep in infants.

The trend of synchronized (high-voltage, low-frequency) electroencephalographic (EEG) activity was determined across nonrapid eye movement (NREM) sleep periods of night sleep in two groups of infants (eight "young infants", age 2-19 weeks; eight "older infants", age 21-54 weeks) and, a reference, in a group of eight adults (age 16-45 years). By automatic analysis of the sleep EEG, a parameter was derived that represents the degree of synchronized (high-amplitude, low-frequency) EEG activity for successive 30-second epochs. For each subject, the average level of EEG synchronization for each NREM period and the time of the NREM period midpoints were determined. In all three groups, synchronized EEG activity tended to decrease across successive NREM periods. This trend was weaker for infants than for adults and, surprisingly, weaker for older than for young infants. This suggests that the decreasing trend typical for adults is already present in the first months after birth. The difference in trend between infants and adults may be caused by differences in the 24-hour sleep-wakefulness distribution, whereas the distinction between young and older infants could be related to a restructuring of sleep in the second semester, in particular to the emergence of slow-wave sleep and its peculiar distribution across NREM periods of night sleep.

Adolescent↗

[The indices of EEG spatial synchronization in normal 10- to 12-year-old schoolchildren and in learning difficulties].

10-12-year-old schoolchildren were divided into three groups: a normal one, a group with psychological problems, and a third one with medical problems. The EEG was recorded at resting and while reading a text. An obviously higher global spatial synchronisation was found in the normal group as compared with the "medical" one. In the "psychological" group, a diffuse hyperresponsiveness was found in the transition state from resting to activity. The above differences seem to be due to a morpho-functional immaturity and a functional insufficiency of two different portions of the brain ascending activating system. Evidently the "medical" group children have an immaturity of the bulbar and midbrain reticular formation, whereas the "psychological" group children have, probably, an immature diencephalic activating system (the thalamus).

Adaptation, Psychological↗

Variability of EEG synchronization during a working memory task in healthy subjects.

Working memory is associated with an increase in EEG theta synchronization and a decrease in lower alpha band synchronization. We investigated whether such changes in mean synchronization level are accompanied by changes in small scale fluctuations of synchronization. EEGs (19 channels; average reference; sample frequency 250 Hz) were recorded in 21 healthy subjects (12 males; mean age 62.5 years; S.D. 2.1) at rest and during a visual working memory condition. EEG synchronization was computed in six frequency bands (2-6; 6-10; 10-14; 14-18; 18-22; 22-50 Hz) using the synchronization likelihood. Variability of the synchronization was quantified with synchronization entropy. During the working memory condition synchronization increased in the 2-6 Hz band, and decreased in the 6-10, 14-18 and 18-22 Hz bands. Working memory was associated with increased variability in the 2-6 Hz band, and decreased variability in the 6-10 Hz band and, to a lesser extent, in the 14-18 and 18-22 Hz bands. Working memory is accompanied not only by characteristic changes in the mean level of interactions between neural networks, but also by changes in small scale fluctuations in such interactions. Strong, but rapidly fluctuating coupling between neural systems might provide a mechanism to optimize the balance between local differentiation and global integration of brain activity.

Adult↗