Neuronal impulse trains of the visual and sensorimotor areas of the rabbit neocortex in calm wakefulness and pseudoconditioning

The conjugation of unit activity in the neocortical visual and sensorimotor areas during calm wakefulness and in intersignal intervals, in two groups of rabbits at pseudoconditioning was studied. The first group was presented in a random order with flashes and electrocutaneous stimuli, the second one--with sounds and electrocutaneous stimuli. The number of neurones pairs working in correlation during calm wakefulness is significantly less (35%) than during pseudoconditioning (49 and 50% in the first and second rabbits groups, respectively). During calm wakefulness and in both groups during pseudoconditioning, the number of pairs with delays of discharges of the visual area neurones after the sensorimotor one, and of the sensorimotor after visual up to 120 ms was equal. Comparison of the data on delayed neuronal discharges during calm wakefulness and pseudoconditioning with those obtained earlier with conditioned reflexes testifies that forestalling of visual area neuronal discharges by sensorimotor discharges is characteristic only for the activity of cortical projections of conditioned and unconditioned stimuli.     

Interaction between neurons of the visual and sensomotor areas of the neocortex during elaboration and extinction of a conditioned defense reflex

Cooperation in activities of pairs of neurones situated in projection cortical areas of conditioned and unconditioned stimuli was studied in rabbits during intertrial intervals at different stages of elaboration and extinction of conditioned defensive response by means of auto- and crosscorrelation analysis of impulse fluxes. At the stage of generalization the number of pairs of neurones discharging in correlation was shown to increase a little (64 per cent) in comparison to that at the initial stage of conditioning (50 per cent) and pseudoconditioning (54 per cent). At the stage of stabilization and during extinction the number of pairs of neurones discharging in correlation decreased correspondingly to 34 and 32 per cent. Parallel analysis of correlation in neuronal discharges and simultaneously recorded electroencephalogram allowed to suppose that excitatory synchronizing influences and inhibitory cortical system play a great part in synchronization of activities of cortical neurones. Participation of these two systems is not the same at different stages of conditioning and extinction.     

Temporal structure of the spike trains of neuronal pairs in the neocortex during calm wakefulness in rabbits

For revealing the conjugation of impulse activity of simultaneously recorded neurones in the visual and sensorimotor areas of rabbits neocortex, histograms were built of cross- and autocorrelation (CCH and ACH) of their discharges sequences. From successions of discharges of the visual cortex neurones, forming pairs of cells acting in correlation, spikes were singled out conjugated with neuronal discharges in the sensorimotor area, and their ACHs were built. In 77% of cases a resemblance was observed of temporal structure of conjugated discharges and real spike sequences, in 23%--there was no such resemblance. Analysis of conjugated discharges ACHs showed that they appeared predominantly periodically (87% of cases). The number of fragments with periodic discharges frequency within delta-frequency range from 3 to 4 Hz prevailed.     

Correlation of the neuronal impulse activity of different areas of the neocortex in rabbits in the switching of positive and inhibitory conditioned reflexes

By cross-correlation method conjugation in impulse activity of pairs of simultaneously recorded units in the visual and sensorimotor areas of the neocortex (representations of the conditioned and unconditioned stimuli) was studied during the switching of defensive positive and inhibitory reflexes in rabbits. 30-second fragments of impulse activity were analyzed in intervals before and after three presentations of positive and inhibitory conditioned stimuli (CS). The activity of single pairs of units in the process of stimuli presentation was characterized by instability. Only in 29% of 51 units' pairs the presence or absence of correlation between their discharges was stable. Analysis of activity of 75 pairs of units showed differences in impulse activity in inhibitory and positive tonic reflexes. The number of pairs of correlated units in about 30 s after inhibitory CS decreased (27%) in comparison with the interval before its action (52%). In the inhibitory tonic conditioned reflex, temporal intervals of discharges of one unit correlated with discharges of the second unit undergo changes. The obtained results testify that such parameter as the number of pairs of units with correlated activity in intersignal time interval is very important for the realization of motor reaction to conditioned stimuli.     

Correlation of neuronal reactions of different regions of the neocortex of the rabbit to conditioned positive and inhibitory stimuli

The activity of neurones of the visual and sensorimotor areas of the neocortex was simultaneously recorded in rabbits under the action of conditioned, inhibitory stimuli (CS, IS) and at simultaneous presentation of CS and the unconditioned stimulus (US) after trans-switching of the positive and inhibitory conditioned reflexes. Mean time of conjugated reactions of simultaneously recorded pairs of neurons is similar under the action of CS, combined action of CS and US and under IS. During the IS action as compared with CS, the phasic activity of some neurones in different areas increased, and due to this the similarity of reactions also increased; in other pairs of neurones the reactions weakened and the responses similarity decreased. The frequency of the appearance of intervals of the conjugated excitation after CS corresponds to 4.3 Hz; under combined action of CS and US it increases up to 5.9 Hz, and after IS--it decreases to 3.6 Hz. The order of coming into conjugated action of different pairs of neurones changes depending on the signal significance of the stimuli.     

The dependence of the neuronal reactions of the sensorimotor cortex to a simultaneous complex stimulus on the level of the differentiation of its components]

On alert animals the change was studied of the neuronal activity of the sensorimotor cortical area of cats brain in dependence on the level of differentiation of the components of simultaneous heteromodal complex stimulus. According to the character of this dependence and a number of other parameters two groups of neurones were singled out in the sensorimotor cortex. It was shown that parameters of reactions of all recorded neurones of the sensorimotor cortex to the positive conditioned signal were the first established after consolidation of the animal conditioned motor activity. In the course of elaboration following parameters changed: expressiveness, intensity, duration and value of latency. Reactions of neurones of both groups to the inhibitory stimuli were stabilized only after consolidation of the habit of differentiation. Responses of the first group neurones changed only by the pattern of discharge, while the responses of the second group neurones could change by expressiveness of response, its sign, duration and value of latency. Oscillations of the differentiation level after finishing of the elaboration of inhibitory conditioned reactions affected only the responses of the second group neurones to complex components.     

Types of correlative relations of the neuronal impulses of the visual and sensomotor areas of the neocortex in the rabbit

Neuronal activity of the visual and sensorimotor cortical areas was simultaneously recorded in intersignal intervals in rabbits with conditioned reflexes to light and sound. ES-1020 computer built cross-correlation (CCH) and autocorrelation (ACH) histograms of impulse activity of the se neurones. Analysis of CCH form, built with a bin of 2 ms gave no convincing proofs of the presence of synaptic connections between neurone of the visual and sensorimotor cortical areas and of influences from a common source with few switchings on interneurones. Analysis of CCH built with bins of 10 and 30 ms, allowed to single out primary and secondary peaks and troughs. Wide primary peaks with the base covering the beginning of the coordinates, were met in 51.9% of cases; wide peaks, greatly shifted relatively to the coordinates beginning,--in 40.9% and trough--in 7.2% of cases. Secondary peaks did not always reproduce the ACH form of impulses trains of recorded neurones. The CCH analysis allowed to suggest two basic mechanisms, eliciting the correlated discharges--action of common input and influence of the neurones of one area on the other through a number of interneurones. It was not always possible to separate these two cases by means of CCH. In a number of cases, a more complicated character of neurones interaction could be suggested.     

Temporal structure of the spike trains in the neuronal impulses of the visual and sensorimotor areas of the rabbit neocortex during conditioned reflex activity

Conjugated spikes were singled out from successions of discharges of neurones in the rabbit's visual and sensorimotor neocortical areas acting in correlation. Their temporal structure was studied at intersignal intervals in transswitching of defensive positive and inhibitory conditioned reflexes and also in pseudoconditioning. The obtained results testify that conjugated discharges appeared for the most part periodically (in the average in 85% of fragments of unit activity). At positive and inhibitory conditioned reflexes, the frequency of periodical conjugated discharges belonged in most cases to the theta-rhythm range. At pseudoconditioning, periodical conjugated discharges were in the main of frequencies of delta-range (up to 4 Hz). The obtained results testify about a great significance of cortical neurones interaction in theta-range frequency at conditioned activity.     

A comparison of the spike train activity of the cortical neurons and of the spatial synchronization of the EEG

In chronic experiments EEG coherence and conjugation of impulse activity were compared of neurones of the visual and sensorimotor areas of rabbits neocortex simultaneously recorded with the same electrodes. Connection was revealed between the presence and properties of conjugated neurones activity and EEG coherence at various frequencies. At correlated neurones activity a greater EEG coherence was observed on frequencies of 3-4,5 Hz than at the independent activity. At the highest level of the EEG coherence the neurones discharged with less delay of one after the other in pairs, and in their synchronization a common source participated more often than at the lowest level of the EEG coherence.     

The characteristics of the neuronal reactions of the cat somatosensory cortex to a heteromodal complex conditioned signal]

On alert cats the change was studied of the activity of the neurones of the sensorimotor cortical area at instrumental reaction to a simultaneous heteromodal complex stimulus. It was shown that in the projection of distal limb areas a group could be singled out of neurones, which changed their activity in one direction depending on the character of presented signals. In these cells an increase of discharges frequency was observed in response to complex stimulus, consisting of light and sound signals. After the extinction of the motor reaction both to the complex stimulus and to its components neuronal reactions of lesser intensity was recorded, what determined the absence of the motor reaction. This group of neurones had receptive fields localized on distal limb areas, it was activated at fulfillment of the movement of catching the reinforcement and belonged to neurones of the pyramidal tract. The neurones with receptive fields on the whole limb surface or changing their activity at the animal pose change, had variable reactions to positive and differentiation stimuli.     

Effect of microelectrophoretic application of GABA on the formation of a cellular analog of a conditioned reaction

Influence of GABA application on ability of sensorimotor cortical neurones to form conditioned cellular reactions was studied in chronic experiments on rabbits. Neuronal reactions were analysed at the initial stage of eye-lid conditioning. GABA application in a period of presentation of paired conditioned and unconditioned stimuli disturbed formation of inhibitory conditioned cellular reactions and did not influence elaboration of activating responses. Plastic changes underlying the process of formation of inhibitory conditioned neuronal reactions are supposed to be of intracortical nature.     

Effect of substance P on behavior in the rabbit and the activity of cortical neurons during training

Subcutaneous injection of substance P to rabbits in a dose of 250 mcg/kg elicited a transitory disappearance of motor reactions to painful reinforcing stimuli and a reduction of their probability to reinforced and inhibitory light flashes, as well as a protracted heart rate increase and decrease of respiration rate. One third of the neurones recorded decreased their background firing level and or excitatory components of the reactions to reinforcement and conditioned light flashes. The decrease was most distinctly seen in the sensorimotor cortex and less pronounced in the visual cortical area and hippocampus. The influence of the substance P on different types of cortical inhibition was not the same. Tonic inhibition of neuronal activity in response to reinforcement was enhanced. Bioelectrical parameters which reflect an enhancement of inhibitory hyperpolarization during elaboration of internal inhibition (i.e. inhibitory firing delays and corresponding background and evoked slow potentials oscillations) were not changed.     

Effect of a derivative of GABA, phenibut, on behavior and the activity of visual cortex neurons of the rabbit during elaboration of a defensive reflex and internal inhibition

In experiments on alert non-immobilized rabbits the effect of subcutaneous administration of the GABA-derivate--phenibut on behaviour, slow potentials and impulse activity of neurones of the visual cortex was studied during elaboration of a defensive reflex to light flashes and of conditioned inhibition. During the action of phenibut late negative-positive components of the evoked potentials to flashes, corresponding inhibitory pauses and postinhibitory activation gradually increased; then stable predominance of slow high-amplitude potential oscillations and corresponding neuronal group bursts appeared, separated by inhibitory pauses and in intersignal periods. Reinforcing stimulus, as before phenibut administration, lowered the amplitudes of slow potential oscillations and weakened inhibitory pauses in neuronal impulse activity. Dynamics of movements in response to the stimuli was of a phasic character. 3 hours after phenibut administration the discrimination of reinforced and inhibitory light flashes has distinctly improved. The obtained results confirm the initial concept of the significant role of the GABA-ergic inhibitory system in the process of elaboration of internal inhibition.     

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.     

Changes in trace reactions of sensorimotor cortex and putamen neurons as affected by haloperidol

Experiments on conscious rabbits were made to elaborate motor conditioned reflexes through pairing stimuli with electrocutaneous reinforcement applied every 30 s. Neuronal activity in the sensorimotor cortex and putamen was recorded during formation and reproduction of the conditioned reflexes before and after haloperidol injection (0.2 mg/kg i. v.). In the putamen, haloperidol increased the number of neurons exhibiting trace conditioned activity and made the intensity and duration of these processes rise. The changes seen in the sensorimotor cortex were opposite in nature. Inhibition of trace conditioned activity in the sensorimotor cortex depended mainly on the decreased amplitude of the reaction conditioned component. The role of the dopaminergic system in the interaction of the neostriatum and sensorimotor cortex and in formation and reproduction of trace conditioned activity of both the structures is discussed.     

The dynamics of the activating and inhibitory types of cortical neuron synchronization during the realization of a defensive reflex and internal inhibition

In the visual and sensorimotor areas of the neocortex and in the hippocampus of alert nonimmobilized rabbits, in response to combinations of light flashes with electrocutaneous limb stimulation an increase was observed of synchronization in the activity of the near-by neurones by activation by inhibitory type (coincidence of the presence and absence of impulse activity). In response to flashes against the light background--conditioned inhibitor--in the visual cortex synchronization of neurones increased by inhibitory type, and in the sensorimotor cortex and hippocampus changes of synchronization appeared, similar to the action of pain reinforcement but considerably weaker. The increase of synchronization by the activation type took place mainly in the neurones pairs with unidirected increase of impulses frequency and by the inhibitory one--with its decrease. Along with this, in a considerable part of neurones pairs both changes of synchronization appeared at the impulses frequency changes of different direction.     

Neurons of the rabbit midbrain reticular formation during a defensive conditioned reflex]

Analysis of unit activity of the midbrain reticular formation was carried out on alert rabbits during defensive conditioning. Most of the examined neurones exhibited phasic responses corresponding in time to the components of the evoked potential (EP) recorded in the cortical visual area in response to the "indifferent" stimulus, and to the conditioned stimulus and electric cutaneous reinforcement. The data obtained are considered from the standpoint of the Anokhin functional systems theory. A conclusion has been made regarding the participation of reticular units in providing all the basic mechanisms of the functional system of the behavioral act. Discharges of one and the same neurone may correspond to different components of the EPs to conditioned and unconditioned stimuli. In different behavioral acts a neurone may apparently participate in different systemic mechanisms.     

Effect of disulfiram on the interconnected activity of cortical neurons in trained cats

The action of disulfiram on interconnected activity of neurones in the visual and motor cortical areas was studied in cats with food-procuring conditioned responses to light. Multiunit activity was recorded from the areas and, by means of amplitude discrimination, separated into impulse flows. Crosscorrelation analysis of the impulse series was used to reveal the character and temporal parameters of interconnected activities of neurones firing in correlation within the limits both of the same cortical area and of the two different ones. A depressing action was shown of the disulfiram on the food-procuring reaction, accompanied by a decrease of the number of pairs of neurones from the visual and motor cortical areas mostly acting in interconnection, interactions with long time delays being mostly affected. The character of action of neighbouring neurones in the visual and motor cortical areas changed in the same direction, expressed in their firing by a "common source" type. The question is discussed of disulfiram influence on interneuronal connections of both types suggesting a decrease of alimentary motivation as well as disturbance of food-procuring conditioned motor coordination.     

Neuronal specialization of the motor cortex in normal rabbits and following destruction of the visual cortex

The activity of neurones of the anterolateral part of the motor cortex in food-acquisition behaviour was compared in two control rabbits and in three rabbits after the operation of bilateral ablation of the striatal cortex. In two of three operated rabbits the pattern of behavioural specialization lost considerably the specificity peculiar to the motor cortex (predominance of G-neurones activated in grasping of food), approaching (but not becoming identical) the pattern of specialization of the visual cortex neurones: the number of G-neurones decreased in a half, and the number of L-neurones (activated in connection with the acts of instrumental food-acquisition behaviour which animals were trained to in the experimental cage) was doubled. Changes of the activity were significantly less expressed in the third operated rabbit. The number of the neurones activated in food-acquisition behaviour in operated rabbits in comparison with the control ones was reduced in the upper layers of the cortex and increased in the lower layers. The resemblance is discussed of the basic processes of animals learning and behaviour recovery.     

Neurophysiologic and biochemical aspects of visual system development in the rabbit under conditions of photic deprivation]

Changes in the functional state of the visual cortex were studied by behavioral and electrophysiological cues and the chemism of its neurones at the cellular and subcellular levels in rabbits raised for one to two months in the dark. It has been shown that visual deprivation leads to retarded dynamics of elaboration and consolidation of the conditioned defensive reflex to light and to changes of opposite signs of the visual cortex surface EPs to specific and non-specific stimuli. Typical of the EPs to photic stimuli is a considerable decrease in amplitude and longer latency as compared with normal, and enhanced amplitude and shorter latency to acoustic stimuli. It has been cytochemically established that about half of the pyramidal neurones of the visual cortex layer V of the experimental animals display features of biochemical underdevelopment (of the size of the cytoplasmatic mass, the protein reserve). Under the same conditions activity depression was revealed in cytochromoxydase, Na, P-ATPhase and ACHE, expressed to a different degree in separate subcellular fractions of the visual cortex. The MAO activity selectively augments in the subfraction of cholinergic synaptosomes. It has been assumed that functional and biochemical changes in different groups of the visual cortex neurones due to deprivation are linked with both the properties of the synaptic structures in regard to perception of impulses of different modalities and the peculiarities of their chemism.