Nonlinear analysis of epileptic activity in rabbit neocortex

We report on the nonlinear analysis of electroencephalogram (EEG) recordings in the rabbit visual cortex. Epileptic seizures were induced by local penicillin application and triggered by visual stimulation. The analysis procedures for nonlinear signals have been developed over the past few years and applied primarily to physical systems. This is an early application to biological systems and the first to EEG data. We find that during epileptic activity, both global and local embedding dimensions are reduced with respect to nonepileptic activity. Interestingly, these values are very low () and do not change between preictal and tonic stages of epileptic activity, also the Lyapunov dimension remains constant. However, between these two stages the manifestations of the local dynamics change quite drastically, as can be seen, e.g., from the shape of the attractors. Furthermore, the largest Lyapunov exponent is reduced by a factor of about two in the second stage and characterizes the difference in dynamics. Thus, the occurrence of clinical symptoms associated with the tonic seizure activity seems to be mainly related to the local dynamics of the nonlinear system. These results thus seem to give a strong indication that the dynamics remains much the same in these stages of behavior, and changes are due to alterations in model parameters and consequent bifurcations of the observed orbits. Received: 5 February 1997 / Accepted in revised form: 18 September 1997     

Suppression of gamma EEG activity as an index of a spreading depression wave in the neocortex of a waking rabbit

A spreading depression (SD) can spontaneously develop in seizures, attacks of migraine, vascular disorders and other pathological states of the brain. However, problems in technique of recording the DC-potential in the neocortex of humans and waking animals substantially restrict the possibilities of studying functional consequences of the SD. In this article, the EEG pattern was studied in detail at the moment of the SD development. Specific features were revealed, which make it possible to detect the SD without recording shifts of the DC-potential. At the moment of the SD arrival, the interhemispheric balance drastically disturbs because of a strong decrease in the high-frequency activity. By the time indices, the course of the suppression of the gammal and gamma2 EEG frequencies is the most reliable symptom of the SD wave development. The EEG spectral power in the delta band increases with a certain delay in reference to the deep depression of the high-frequency activity and is, in essence, an SD aftereffect. The found EEG signs of an SD wave can substantially simplify the identification of this phenomenon both in experiment and clinical conditions in certain pathological states of the brain.     

Spreading depression in a waking rabbit with the spectral EEG analysis

It was shown that the manifestation of the SD phenomenon in dynamics of the cortical high-frequency gamma activity is rather prominent after bipolar interpretation of the common reference electrode derivations, i.e. when the modeling of the bipolar signal on the base of monopolar EEG recordings is used. The SD wave was accompanied by significant decrease in the power of the EEG gamma band (37-47 Hz) in all observed cortical areas. A curve of decline of gamma activity power had distinct fore and back fronts, so the time of SD appearance in different cortex areas and it's spread succession could be well determined. In the long-term experiments SD waves were characterized by normal (i.e. successive) spread through the cortex mainly in initial three-four experiments. In the subsequent tests latency of SD waves in different cortex areas changed and disturbance of propagation became obvious. Sometimes SD arose rapidly (due 0.5-2 min) on the whole dorsal neocortical surface, when the standard injection of the KCI was done. In the most of experiments the delay of the SD wave appearance was prolonged till 6-9 min or no SD wave occur at some cortical regions. Week epileptiform activity could conduct abnormalities in the SD. In many instances electrophysiological signs of the cortical excitability changes were absent. However the modified spatial SD characteristic and spontaneous occurrence of the repeated. SD waves indicated the increased functional inhomogeneities of the neighboring cortical areas. So, spectral EEG analysis in awake rabbits made it possible to characterize the SD wave both in case of its normal propagation through the cortex and in unusual forms of this reaction.     

Structural organization of rabbit neocortex in prenatal ontogenesis

In the review on the basis of data of different authors, the development dynamics of the rabbit neocortex in prenatal life is presented. The main points of neurons development, interneuronal contacts, layer-by-layer and areal differentiation as well as neocortex afferental and efferental communications, that are elucidated ambiguously by investigators, are discussed. For the purpose of determination of the neocortex maturation to the birth-date the comparative analysis of the reference morphological data and author's own neurophysiological data is accomplished.     

A spectral-correlational analysis of the electrical activity of the sensorimotor cortex and the internal geniculate body during a motor-polarization dominant]

By spectral-correlative analysis the dynamics was studied of structural changes of coherent relations of the electrical activity of the sensorimotor cortex and medial geniculate body (MGB) of the rabbit under motor polarization dominant created by the action of DC anode on the sensorimotor cortex area. It was shown, that in the power spectra of the MGB, besides the increase of the delta-region components in interstimulus intervals at the dominant, during the action of the sound stimulus a distinct maximum appeared in the alpha-rhythm range of the electrical activity of MGB of the "dominant" hemisphere. The coherence increase of the delta-range of the MGB and the sensorimotor cortex electrical activity in the "dominant" part of the brain was manifested exclusively in the period of sound stimulus action.     

Quantal analysis of synaptic processes in the neocortex

The application of fluctuation analysis to studies of synaptic function in the neocortex is discussed. Analysis of failures of transmission has been valuable in indicating whether a presynaptic or a postsynaptic site is responsible for a change in synaptic efficacy. When combined with detailed ultrastructural verification of all synapses involved in an individual cell to cell connection, a reasonable estimate of quantal size and release probability under conditions of low frequency activity can be obtained. However, both the number of available release sites in functional terms and the probability that an action potential (AP) will release transmitter from any given site can vary from AP to AP at higher frequencies. A variety of presynaptic mechanisms that modulate release are now apparent. For example, one mechanism dominates release patterns at one class of connection which is insensitive to absolute firing frequency, but responsive to changes in frequency. At another class of connection, a different mechanism dominates, resulting in high sensitivity to frequency.     

The spectral correlation characteristics of the brain electrical activity in the rabbit during thirst]

Spectral-correlation analysis of the summate electrical activity of a number of subcortical structures of rabbit brain, having, by literature data, a relation to drinking behaviour showed that the increase of drinking excitability, induced by water deprivation was accompanied by definite reconstruction of biopotentials. In electrical activity of the studied structures, the spectrum power, as a rule, decreased (except in the paraventricular nucleus), and a definite structure of coherent connections between the subcortical and cortical potentials was established. Among the studied subcortical formations, structures (anterior hypothalamic area, lateral preoptic area, medial preoptic area, paraventricular nucleus) could be singled out where reconstructions of spectral characteristics of biopotentials took place most regularity, and the electrical processes in which were characterised by coherence index by an increase of spatial interconnection with the neocortex potentials, what allows to consider them as most significant for organization of drinking excitation.     

EEG and behavioral effects of gamma-hydroxybutyrate in the rabbit

The effects of gamma-hydroxybutyrate (GHB) upon sleep wakefulness patterns and quantified nuchal muscle activity were examined in the rabbit in a dose-response paradigm (25–1,000 mg/kg). Relative to control (saline) values, there was no facilitation of sleep onset or epileptogenic activity at any of the dosages studied. However, at the higher GHB concentrations, slow wave sleep and tonic muscle activity were enhanced and a high amplitude, slow activity was superimposed on background EEG patterns. The highest concentration of GHB (1,000 mg/kg) was associated with depression of motor activity. An enhancement of paradoxical sleep observed at lower GHB levels in other species occured in attenuated form in the rabbit. The results indicate dose-related effects on both sleep and motor activation in the rabbit, but the absence of seizure activity for the concentrations of GHB studied.     

Coherent analysis of the EEG during development of trace processes of the polarization dominant in the rabbit

By the method of spectral-correlation analysis, the dynamics was studied of changes in intercentral relations of electrical activity in the sensorimotor and premotor cortices of both cerebral hemispheres during formation, course, recovery and extinction of motor dominant created by a single action of DC anode in rabbits sensorimotor cortex. The motor dominant is capable to be preserved during the test for a long time after DC switching off and to recover in subsequent days under the action of testing stimuli. It should be noted that the recovery of the structure of intercentral relations of electrical brain activity, characteristic of dominant state, takes place much earlier than the motor "dominant" reaction.     

Electrophysiologic analysis of the conditioned reflex activity of cats against a background of thirst

EEG of the cerebral cortex and hypothalamic electrogram of cats during salt load and water deprivation were recorded at performance of conditioned running elicited by signals of insipid and salt food. The signal of insipid food presented during thirst was accompanied only by the cerebral cortex activation, while the salt signal involved in activation not only the cortex but the hypothalamus as well. In cases when the insipid signal was reinforced by salt food and the animal ate it (though during thirst it rejected the food), strong cortex activation was observed with the involvement of paraventricular parts of the hypothalamus. During intense thirst reversal was obtained of the signal role of the conditioned cue in accordance with the new quality of the alimentary reinforcement. Hypothalamo-cortical mechanisms of dominant motivation and its conditioned provision are discussed.     

Segmentation of the epileptic EEG by means of a finite state automation

The aims and merits of the syntactical decomposition of the epileptic EEG are discussed in some detail. General assumptions in the design of segmenting algorithm - an essential part of the reduction of data program - are considered and a working example of segmenting routine os presented. This routine is designed as a finite state automaton, equivalent to a formal regular grammar. The performance of the routine on two sets of experimental data is presented and its intrinsic stability is shown in terms of cluster analysis of the segments produced. Means of extension of the proposed scheme to other cases of potential interest as well as possible applications in the higher level syntactical analysis are pointed out.     

Long-term changes in EEG spectra of the hippocampus and neocortex during pharmacological action on the cholinergic system

Statistical analysis of EEG spectra averaged over 10-min periods showed that inhibitor of acetylcholinesterase physostigmine induced the long-term (tens of minutes) characteristic changes in the electric activity of the dorsal hippocampus (CA1 field) and somatosensory cortex of unrestrained rats. With increasing the physostigmine dose from 0.05 to 0.5 or 1 mg/kg the averaged power of the theta-rhythm did not rise in the range of 3.6-4.9 Hz and was suppressed in the range of 5.7-11.9 Hz both in the hippocampus and neocortex. The maximal frequency shifted to the left (from 3.6-6.4 to 3.6-4.9 Hz). In contrast to this, the averaged power in the delta (1-1.5 Hz)-I and beta-2 ranges (20.3-26.5 Hz) significantly nonlinearly increased and that of the beta-1 substantially decreased. Scopolamine eliminated all extrema of the hippocampal and neocortical EEG spectra induced by physostigmine, which is indicative of the role of M-cholinoreceptors in these effects. The increased dose of physostigmine (1 mg/kg) produced inversion of the ratio between the averaged power of beta-2 in neocortex and hippocampus: it became significantly higher than in the neocortex. All these data suggest that the mechanisms of cholinergic modulation of the theta- and beta-rhythms are essentially different. We think that significant enhancement of the content of endogenous acetylcholine content produce a long-term tonic decay of the functional activity of the hippocampus and neocortex and play an important role in the mechanisms of dissociated states of memory and consciousness, contextual learning and conditioned switching.     

Analysis of the theta-rhythm of the rabbit neocortex by estimation of particular coherence functions

Situations characterized by differences in the genesis of the theta-rhythm were distinguished by the analysis of activity recorded simultaneously from various points of the rabbit neocortex and hippocampus by means of the estimation of particular coherence functions. With maximal cross-correlation between potentials from different parts of the neocortex, in most cases (not less than 75%) the high coherence of these potentials in the theta-band is determined by a theta-rhythm generated actually in the neocortex. In about 20% of cases the high coherence of these potentials may be the result of the passive conduction of hippocampal theta-activity to the surface of the neocortex. To increase the likelihood of the passage of excitation from the afferent to the effector system active generation of rhythmic waves in the theta-band in both the neocortex and hippocampus is essential. Rhythmic processes in both structures must be coherent with each other.     

Temporal patterns analysis of spontaneous unit activity in the neocortex

An analysis has been made of the spontaneous firing of neurons from the posterior motor cortex of the rabbit. Extracellular recordings were made by means of glass capillary micro-electrodes. The recorded neural activity was processed with a general purpose computer and an adapted data processing system. The impulse interval series were analysed by means of a statistical time series analysis. In the first part of the paper, the first order properties of the neutral activity are examined. Using a mathematical model for data reduction, an overall picture of the first order properties of the activity of the set of recorded neurons are obtained. In the second part of the paper, the second order properties of this activity are examined. Using another simple mathematical model completing the first one, the principal aspect of the second order properties are summarized.