Analysis of theta-rhythms recorded in an isolated strip of rabbit sensomotor cortex

To test the possibility that the theta-rhythm may spread passively from the hippocampus into the neocortex in rabbits and to analyze the properties of these rhythms recorded from the surface of the neocortex, an investigation was made of activity recorded simultaneously from the hippocampus and various points of the neocortex, including from a strip of the sensomotor cortex isolated completely with respect to its nervous connections. The use of special correlation methods showed that an appreciable contribution to the potentials recorded from the isolated strip can be made by the passive conduction of hippocampal activity (provided that it generates a regular high-amplitude theta-rhythm). At the same time, potentials in a theta-rhythm may be generated in the isolated strip itself. The theta-rhythm recorded in the intact rabbit cortex may thus be of combined genesis: generated in the neocortex itself and spreading passively from the hippocampus through the brain tissues as through a volume conductor.     

An investigation of phase relations between theta-rhythm potentials in the neocortex and hippocampus of rabbits

Phase relations between rhythmic activity recorded from the hippocampus and from various levels of the neocortex were calculated on a two-dimensional model on the assumption of the passive spread of hippocampal activity. The calculation showed that if such a spread of activity takes place an appreciable phase shift, caused by the spatio-temporal structure of the hippocampal source of the theta-rhythm, may exist between the rhythmic components of potentials recorded from the neocortex and hippocampus. In some series of electrophysiological experiments on rabbits various consequences of the model calculations were tested. Phase relations of the theta-rhythm recorded in the neocortex and in the hippocampus in the presence of maximal correlation between cortical potentials cannot be explained, in the overwhelming majority of cases, purely by the passive spread of hippocampal activity to the surface of the neocortex.     

Dependence of the spectrum of electrical activity of the neocortex and hippocampus in rabbits on the intensity of the stimulation of the midbrain reticular formation

Study of dominating spectral maxima in delta-, theta- and alpha-ranges of the electrical activity of rabbits' neocortex and hippocampus showed that an increase of the frequency of the mesencephalic reticular formation stimulation from 60 to 200 imp/s led in both structures to an enhancement of the theta-rhythm (up to 130% in the neocortex and 147% in the hippocampus) and suppression of delta- and alpha-activity (correspondingly up to 67 and 34% in the neocortex and 37 and 48% in the hippocampus) with subsequent weakening of this effect at frequency increase up to 1000 imp/s. In the hippocampus, the reticular stimulation was more effective with respect to the theta- and delta-rhythms, and in the neocortex--with respect to the alpha-rhythm. In both structures the theta-rhythm amplitude changed less than the amplitude of the delta- and alpha-activities. Dependence of the amplitude of dominating rhythms on intensity of reticular formation stimulation differed from the analogous frequency dependence of the same rhythms.     

Correlation between bioelectrical processes of the cortex, thalamus, midbrain reticular formation during formation of a defensive conditioned reflex in rabbits]

EEG power spectra of the sensorimotor area of the neocortex, the dorsal hippocampus, midbrain reticular formation and anteroventral thalamic nucleus, as well as corresponding coherence functions and phase spectra, undergo changes during formation and performance of defensive conditioned reflex in rabbit. The conclusion is draen that in the process of conditioning a morphofunctional system of brain structures is established including the above mentioned formations. Their functional integration occurs on the basis of theta-rhythm. The execution of a conditioned act requires isorhythmicity of electrical processes within the theta-range in the studied structures and an adequate level of their excitability, which is manifested in the dominance of 6,0 c/s frequency.     

Spectral correlation parameters of neocortical potentials in the rabbit undergoing paired isorhythmic stimulation

At pairing of isorhythmic stimuli beyond the theta-rhythm frequency limits (3 and 8 Hz), in power spectra of EEGs of the sensorimotor and visual neocortical areas of rabbits, the frequencies are present both of the theta-range and of the stimulation frequency, in the background activity as well as during the stimulation. Both rhythms are in reciprocal relations. The frequency of the theta-rhythm approaches the frequency divisible by that of the stimulation. Under the action of the conditioned stimulus, crosscorrelation coefficients (CC) between the potentials of the areas under study decrease in most cases in comparison to their background values. Combination of the conditioned stimulus with the unconditioned one, leads approximately in equal number of cases to an increase or decrease of CC. After elimination of the stimuli, in most cases CC increases. CC of the background activity does not increase in the course of paired stimuli presentation though a conditioned response is being formed. At presence of stimuli frequency fluctuations simultaneously in the potentials of both areas, the rise of coherence function at this frequency does not occurs always. Thus, the above spectral-correlation parameters of rabbit's cortical potentials differ from those which arise at pairing of continuous nonrhythmic stimuli. This difference is probably due to different characteristics of the stimuli presented.     

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.     

Changes in the bioelectrical activity and temperature of paired brain formations in the rabbit in a single ethanol administration

In chronic experiments on rabbits, in initial period of single action of ethanol, reduction was observed of the dominating frequency of the theta-rhythm and synchroneity of summated bioelectrical activity of homotopic zones and suppression of EEG manifestation but preservation of parameters of the interhemispheric zeta-wave and of temperature asymmetry in the dorsal hippocampus in response to electro-cutaneous stimulation. Late period of ethanol action was characterized by lowering of the amplitude of superslow activity, its inversion in symmetrical areas of the visual neocortex and instability of other parameters.     

Oscillatory models of the hippocampus: A study of spatio-temporal patterns of neural activity

Spatial patterns of theta-rhythm activity in oscillatory models of the hippocampus are studied here using canonical models for both Hodgkin's class-1 and class-2 excitable neuronal systems. Dynamics of these models are studied in both the frequency domain, to determine phase-locking patterns, and in the time domain, to determine the amplitude responses resulting from phase-locking patterns. Computer simulations presented here demonstrate that phase deviations (timings) between inputs from the medial septum and the entorhinal cortex can create spatial patterns of theta-rhythm phase-locking. In this way, we show that the timing of inputs (not only their frequencies alone) can encode specific patterns of theta-rhythm activity. This study suggests new experiments to determine temporal and spatial synchronization. Received: 31 July 1998 /Accepted in revised form: 20 April 1999     

Directed Communication between Nucleus Accumbens and Neocortex in Humans Is Differentially Supported by Synchronization in the Theta and Alpha Band

Here, we report evidence for oscillatory bi-directional interactions between the nucleus accumbens and the neocortex in humans. Six patients performed a demanding covert visual attention task while we simultaneously recorded brain activity from deep-brain electrodes implanted in the nucleus accumbens and the surface electroencephalogram (EEG). Both theta and alpha oscillations were strongly coherent with the frontal and parietal EEG during the task. Theta-band coherence increased during processing of the visual stimuli. Granger causality analysis revealed that the nucleus accumbens was communicating with the neocortex primarily in the theta-band, while the cortex was communicating the nucleus accumbens in the alpha-band. These data are consistent with a model, in which theta- and alpha-band oscillations serve dissociable roles: Prior to stimulus processing, the cortex might suppress ongoing processing in the nucleus accumbens by modulating alpha-band activity. Subsequently, upon stimulus presentation, theta oscillations might facilitate the active exchange of stimulus information from the nucleus accumbens to the cortex.     

Coherent-phase characteristics of the cortical potentials over a wide frequency range in dogs in a state of calm wakefulness

The spectra of coherence (Coh) and phasic shifts (PhS) of cortical potentials were studied by correlation-spectral analysis in a wide frequency range (1--100 Hz) in dogs in a state of quiet wakefulness. Estimation was made by parameters: significant Coh (above 0.5), high Coh (above 0.75), considered PhS (not above +/- 30 degrees), small PhS (not above +/- 15 degrees C). The presence is shown of individual features in coherent-phasic characteristics of neocortex potentials. High values of cross-correlation coefficients (Cc) between cortical potentials correspond in Coh spectra to a great part (90--100% peaks) of significant Coh, in which the great part (90--70% peaks) belongs to the high Coh with small PhS (in most cases). A certain correspondence, though expressed weaker, persisted also for lower values of Cc, Coh, and considered PhS. Coherent-phasic relations between potentials of certain cortical areas are characterized by stability in each animal; a tendency to it is manifest in all animals, despite individual differences of the brain electrical activity.     

Hippocampal memory consolidation during sleep: a comparison of mammals and birds

The transition from wakefulness to sleep is marked by pronounced changes in brain activity. The brain rhythms that characterize the two main types of mammalian sleep, slow‐wave sleep (SWS) and rapid eye movement (REM) sleep, are thought to be involved in the functions of sleep. In particular, recent theories suggest that the synchronous slow‐oscillation of neocortical neuronal membrane potentials, the defining feature of SWS, is involved in processing information acquired during wakefulness. According to the Standard Model of memory consolidation, during wakefulness the hippocampus receives input from neocortical regions involved in the initial encoding of an experience and binds this information into a coherent memory trace that is then transferred to the neocortex during SWS where it is stored and integrated within preexisting memory traces. Evidence suggests that this process selectively involves direct connections from the hippocampus to the prefrontal cortex (PFC), a multimodal, high‐order association region implicated in coordinating the storage and recall of remote memories in the neocortex. The slow‐oscillation is thought to orchestrate the transfer of information from the hippocampus by temporally coupling hippocampal sharp‐wave/ripples (SWRs) and thalamocortical spindles. SWRs are synchronous bursts of hippocampal activity, during which waking neuronal firing patterns are reactivated in the hippocampus and neocortex in a coordinated manner. Thalamocortical spindles are brief 7–14 Hz oscillations that may facilitate the encoding of information reactivated during SWRs. By temporally coupling the readout of information from the hippocampus with conditions conducive to encoding in the neocortex, the slow‐oscillation is thought to mediate the transfer of information from the hippocampus to the neocortex. Although several lines of evidence are consistent with this function for mammalian SWS, it is unclear whether SWS serves a similar function in birds, the only taxonomic group other than mammals to exhibit SWS and REM sleep. Based on our review of research on avian sleep, neuroanatomy, and memory, although involved in some forms of memory consolidation, avian sleep does not appear to be involved in transferring hippocampal memories to other brain regions. Despite exhibiting the slow‐oscillation, SWRs and spindles have not been found in birds. Moreover, although birds independently evolved a brain region—the caudolateral nidopallium (NCL)—involved in performing high‐order cognitive functions similar to those performed by the PFC, direct connections between the NCL and hippocampus have not been found in birds, and evidence for the transfer of information from the hippocampus to the NCL or other extra‐hippocampal regions is lacking. Although based on the absence of evidence for various traits, collectively, these findings suggest that unlike mammalian SWS, avian SWS may not be involved in transferring memories from the hippocampus. Furthermore, it suggests that the slow‐oscillation, the defining feature of mammalian and avian SWS, may serve a more general function independent of that related to coordinating the transfer of information from the hippocampus to the PFC in mammals. Given that SWS is homeostatically regulated (a process intimately related to the slow‐oscillation) in mammals and birds, functional hypotheses linked to this process may apply to both taxonomic groups.     

Functional differentiation of the theta-rhythm of the hippocampus in the cat

On the model of discriminating alimentary conditioned reflex with spatial separation of the sources of conditioned (CR, light flashes) and unconditioned (feeding trough) stimuli the theta-rhythm of the dorsal hippocampus was evaluated in cats. Two types of the theta-rhythm were observed in the spectrum of the hippocampal electrical activity: low-amplitude, consisting of a slow growing theta-waves (type I) and high-amplitude consisting of rapidly increasing theta-waves (type II). The type I theta-rhythm is sensitive to noradrenaline and correlates with behavioural forms directed immediately to the realization of alimentary motivation, while the type II is sensitive to serotonin and correlates with behavioural forms directed to the source of the conditioned stimulus. Enhancement of the type II theta-rhythm takes place during a delay of the expected conditioned stimulus. It is suggested that type I theta-rhythm reflects a level of activity of the brain structures connected with unconditioned mechanisms, with realization of biological motivations, while type II theta-rhythm is connected with conditioned reinforcing stimuli; it correlates with various forms of conditioned orienting reflexes and reflects the work of the "nervous model of reinforcing stimulus".     

The cross-correlation relations of the electrical activity of the cortical structures at different levels of the activation of the midbrain reticular formation in the rabbit]

Changes of correlation coefficients and coherence of the delta-, theta- and alpha-rhythms were studied between the somatosensory, motor and visual neocortex areas, dorsal hippocampus and dentate fascia at stimulation frequencies of the midbrain reticular formation from 60 to 1000 imp/s. It was shown that correlation coefficient between the structures studied increased at 60-200 imp/s and decreased at the further increase of stimulation frequency. In pairs with the visual area the correlation coefficient changed but little. The delta-rhythms coherence tended to decrease with the increase of stimulation frequency. Coherence of the theta-rhythms between the neocortical areas and the hippocampus increased with the increase of stimulation frequency up to 200 imp/s and decreased at higher frequency stimulation while in pairs of these areas with the dentate fascia it continued to rise with the increase of stimulation frequency up to 1000 imp/s. The coherence of the alpha-rhythms was almost unchanged at 60-200 imp/s and mostly had an increase at higher frequency stimulation.     

The intercentral relations of the frontal cortex and limbic structures of the brain in dogs]

In 3 dogs with implanted electrodes, in conditioned experiments correlation of the bioelectrical processes was studied by coherence function calculation of the hippocampus, hypothalamus, amygdala and frontal cortex biopotentials. It was shown, that the level of maximum values of coherence function of bioelectrical oscillations, led from various pairs of the studied brain structures significantly differed both in magnitude and frequency at which the greatest synchronization of biopotentials was noticed. In one dog with a high degree of connection between the hippocampus and hypothalamus biopotentials oscillations, a low synchronization of the frontal cortex and amygdala oscillations was found; in two other animals with a higher level of coherence between the oscillations of the frontal cortex and amygdala biopotentials, a lower degree of connection between the oscillations led from the hippocampus and hypothalamus was revealed. Synchronization of the biopotentials of the hippocampus and frontal cortex and also of the hippocampus and amygdala biopotentials proved to be low in all experimental dogs, what additionally testifies to different role of these structures in organization of the behaviour.     

The effect of hypothermia on the rat radioresistance

The cooling of Wistar rats up to 15-19 degrees C under a condition hypoxia-hypercapnia increased the radioresistance with a dose reduction factor (DRF) of 1.4. To elucidate the mechanisms of hypothermia radioprotective effect was evaluated the functional state of rat neocortex using a electroencephalogram (EEG) as well as was studied the lipid composition of neocortex under the conditions of both normothermia and hypothermia. At 19-20 degrees C the activity within a wide range of frequencies in EEG was suppressed; the nonregular slow waves were recorded against a background of "silence". The reduction of EEG spectrum with increasing temperature began with the low frequencies. At 26-28 egresC the contribution of theta-rhythm (an indicator of brain activity level) in EEG reaches the normothermia value, from this point the rat brain starts to functionate as a whole system. At normothermia the similarity of neocortex lipid composition in nonhibernators (rats) and hibernators (ground squirrels) mammalians was noted. The difference is only in a higher content of phosphatidylinositol in rats. Rats falling into hypothermia state as well as ground squirrels into torpor is followed by a decrease of cholesterol content and the absence of significant changes of the phospholipid composition in neocortex tissues.     

Theta oscillations by synaptic excitation in a neocortical circuit model

Neocortical theta-band oscillatory activity is associated with cognitive tasks involving learning and memory. This oscillatory activity is proposed to originate from the synchronization of interconnected layer V intrinsic bursting (IB) neurons by recurrent excitation. To test this hypothesis, a sparsely connected spiking circuit model based on empirical data was simulated using Hodgkin-Huxley-type bursting neurons and use-dependent depressing synaptic connections. In response to a heterogeneous tonic current stimulus, the model generated coherent and robust oscillatory activity throughout the theta-band (4-12 Hz). These oscillations were not, however, self-sustaining without a driving current, and not dependent on N-methyl-D-aspartate receptor synaptic currents. At realistic connection strengths, synaptic depression was necessary to avoid instability and expanded the basin of attraction for theta oscillations by controlling the gain of recurrent excitation. These results support the hypothesis that IB neuron networks can generate robust and coherent theta-band oscillations in neocortex.     

Cortico-hippocampal relation of electrical activity during a motor polarization dominant in rabbits

By the method of spectral-coherent analysis, the intercentral relations were studied of the electrical activity of the sensorimotor and premotor cortices and of CAs field of the dorsal hippocampus of both hemispheres during the motor polarization dominant, created by the action of the direct current on the rabbits sensorimotor cortical area. The formation was shown of a new structure of the intercentral relations of electrical activity of the sensorimotor cortex and CA3 of the dorsal hippocampus. It should be noted that the dominant optimum and its inhibition are characterized by different interhippocampal relations: at the optimum a low range of the theta-rhythm is highly coherent while at the activation of "the non-dominant" hemisphere--a high range.     

Emotional stress during conditioned reflex transformation of heterogeneous instrumental reflexes

The factors leading to pronounced emotional stress accompanied by changes in electrical activity of hippocampus were studied in four dogs during elaboration of conditioned switching-over of heterogeneous instrumental reflexes. Objectively recorded signs of emotional stress were observed in two cases: when the animal was not accurate enough in discerning defensive and alimentary situations and throughout the improvement of instrumental defensive reflex. Pronouncement of hippocampal theta-rhythm increased at those stages of switching-over elaboration where the probability of successful avoiding pain stimulation was relatively low. The data obtained permit the conclussion that theta-rhythm is the correlate of the degree of the emotional stress and not only a manifestation of general arousal reaction depending on the level of sensory input.     

Traveling Theta Waves along the Entire Septotemporal Axis of the Hippocampus

A topographical relationship exists between the hippocampus-entorhinal cortex and the neocortex. However, it is not known how these anatomical connections are utilized during information exchange and behavior. We recorded theta oscillations along the entire extent of the septotemporal axis of the hippocampal CA1 pyramidal layer. While the frequency of theta oscillation remained same along the entire long axis, the amplitude and coherence between recording sites decreased from dorsal to ventral hippocampus (VH). Theta phase shifted monotonically with distance along the longitudinal axis, reaching ～180° between the septal and temporal poles. The majority of concurrently recorded units were phase-locked to the local field theta at all dorsoventral segments. The power of VH theta had only a weak correlation with locomotion velocity, and its amplitude varied largely independently from theta in the dorsal part. Thus, theta oscillations can temporally combine or segregate neocortical representations along the septotemporal axis of the hippocampus.     

Subcellular changes in the cortex of the motor analyzer and hippocampus of dogs with locally manifested neuroses

In experiments on dogs with local neurosis-continuous flexion of the foreleg-changes were revealed in the beta-rhythm amplitude and the frequency of mean unit activity in the motor cortex, and the appearance and increased amplitude of the theta-rhythm in the hippocampus. Specific activity of Na+-K+-activated, and Mg2+-dependent ATPase decreases in subcortical fractions of the experimental animals' cerebral cortex by 55.0% in the synaptic membranes and 2 to 2.5 times in light and heavy synaptosomes, respectively. In similar fractions of the dorsal hippocampus, the activity of the enzyme decreases by 30.0% in the synaptic membranes and increases by 16.6% in the light synaptosomes and by 6.6% in the heavy ones.