Analysis of changes in the electrical activity of the neocortex in dogs during the formation of the stereotype of food-procuring conditioned reflexes in them

A system of food-procuring conditioned reflexes (dynamic stereotype after I.P. Pavlov) was elaborated in dogs. In the interstimuli periods, 0.6 to 0.8 sec. prior the action of the conditioned stimulus, they exhibited a "state of expectancy" characterized by an increase of frequency (up to 80 per sec.) of potential oscillations of a small amplitude (20 to 30 mcv) and their pronounced sychroneity, predominantly in the anterior parts of the neocortex. The conditioned signal acting against such background produced specific reactions in the form of a limited number of bursts (3 to 4 in 0.5 sec.) of highfrequency (up to 100 per sec.) synchronized activity (HSA) of considerable amplitude (50 to 60 mcv) which always preceded conditioned reactions, being in certain temporal relations with them. A correlation-spectral analysis has shown that in the HSA period, electrical activity became considerably more regular; in the intensity spectra the extreme frequencies of the analyzed band (5 to 7 c/s and 90 to 100 c/s) became prominent, and in most cases high values of the coherence function were due to them. HSA reactions are regarded as a major link in the trigger mechanism of conditioned food-procuring reactions.     

Effect of the type of reinforcement on conditioned reflex activity in male and female white rats

On 24 males and 26 females of white rats of a mixed line, a study was carried out of the speed of formation and consolidation of instrumental drinking and defensive conditioned reflexes, elaboration of differentiation, changes in conditioned activity by the use of stochastic stereotype and by changing the meanings of the conditioned signals to opposite. Conditioned reflexes were elaborated more rapidly in female rats at drinking reinforcement and in male rats--at the defensive one. Differentiations were elaborated more rapidly and were better expressed in females than in males at both types of reinforcement.     

Formation of combined stereotypes of systems of secretory conditioned reflexes

A rhythmic stereotype, followed by a rhythmic-mozaic one of positive and inhibitory conditioned secretory reflexes were successively elaborated in dogs with stabile stereotype of three heteromodal randomely organized systems. Thus a combined stereotype was formed of five randomely and stereotypically organized systems of conditioned reflexes, achieved at intervals of 24 hours or longer in a regular sequence. It has been shown that in the course of formation of the combined stereotype the general functional state (tone) of the cerebral cortex is enhanced. The preliminarily formed specialized systemic nervous process with a similar structure is of major importance for the rate of formation of the complex rhythmic-mozaic stereotype.     

Effect of streptomycin sulfate on the process of higher nervous activity recovery in dogs in the postanesthesia period]

The effect of streptomycin on recovery of the higher nervous activity during the postanesthesia period was studied on 3 dogs with the method of the conditioned reflexes. It was found that streptomycin sulphate administered in a dose of 25 mg per 1 kg body weight induced no changes in the conditioned reflexes of the non-anesthetized animals. After ether anesthesia disorders in the conditioned and unconditioned reflexes, as well as differentiation was observed during the postanesthesia period. On an average they normalized in 7, 6 and 2 days respectively from the moment of anesthesia. When streptomycin sulphate was used during the post anesthesia period, both the strength and the character of the disorders in the conditioned reflexes changed. In this case recovery of the conditioned and unconditioned reflexes was registered in 2 days, while that of the differential inhibition was registered in 3--7 days from the moment of anesthesia.     

Systems organization of cerebral processes in the cat during learning while undergoing electrical stimulation of the neocortex

A delayed spatial choice (DSC) was elaborated in five cats during electrical stimulation (ES) of the frontal cortical parts with a frequency of 80 imp/s. Then the task fulfillment was tested following cessation of the electrostimulation or under the action of other current frequencies (35,3 and 120 imp/s). The number of erroneous choices increased in the absence of ES or during 3 imp/s ES of the frontal parts. The effects of frequencies of 35,80 and 120 imp/s did not differ. Three cats were trained to DSC without ES. The conditioned reflexes were tested during ES of the frontal cortical parts with current frequency of 3 and 80 imp/s. Statistically significant increase of the number of erroneous responses took place in both situations. The obtained data are discussed from the point of view of the integrative activity of the cerebral structures in DSC. Its disturbance during ES of the cortical zones and dissociation (discordance) after cancellation of stimulation during which the conditioned reflex had been elaborated, point to systemic organization of brain functioning in goal-directed forms of behaviour.     

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.     

Typological characteristics of conditioned reflex learning in monkeys

Three types of conditioned learning dynamics were singled out in males of the rhesus monkeys by the speed of formation of a positive motor instrumental reflex and differentiation. Type I is characterized by the fastest mastering of separate subprograms and of the whole test program of conditioned behaviour, by optimal functional characteristics in the process of learning and by a high adaptivity in new conditions. The most prolonged formation of the conditioned reflexes occurs in monkeys of the III-d type; 50% of them do not master the whole learning program; monkeys with such type of learning dynamics have no adaptation of previously acquired conditioned stereotype to new environmental conditions. Monkeys of the II-nd type of learning dynamics possess intermediate characteristics. A notion is substantiated of interconnection of higher nervous activity properties with typological characteristics of conditioned learning dynamics in monkeys.     

Excitability cycles of direct cortical responses during elaboration of conditioned defense reflexes in dogs]

Direct cortical responses (DCRs) to paired stimuli were studied in chronic experiments in dogs during elaboration of classical and instrumental defensive conditioned reflexes. The DCRs were recorded with 20 to 250 ms intervals between stimuli. Paired and single electrical stimulations of the middle suprasylvian gyrus given with a frequency of one per second were used as conditioned stimuli and were reinforced in a similar way. During electrical cutaneous stimulation of the dog's paw and to an even greater extent during isolated action of the conditioned stimulus the initial negativity of the testing DCR became shorter and the degree of its depression diminished. In the case of a following period of facilitation, its degree became greater. It was higher at a distance of 4 to 5 mm from the point of stimulation than at a distance of 2 to 3 mm. During isolated action of the conditioned stimulus, the degree of facilitation was higher than at the period of the possible action of the unconditioned stimulus. The greatest shorterning of the DCR excitability cycle was observed immediately before and during the conditioned lifting of the dog's paw. Excitability cycles of DCR, and possibly of other evoked potentials as well, are a more sensitive indicator of the function state of the cerebral cortex than responses to single stimuli. For this reason it appears promising to use them in studying conditioned reflexes.     

Changes in conditioned reflex activity during adaptation of the rat to pressure-chamber and high-altitude hypoxia

A group of rats with a stereotype of conditioned reflexes was preliminarily trained to hypoxia effects during 30 days (at the "altitude" of 6000 m, time of exhibition--from 10 to 60 min, for 18 days--only 60 min). Adaptive changes in the process of training consisted in a weakening of differentiation inhibition, partial amnesia of the conditioned reaction of active avoidance and appearance of phasic states (equalization and paradoxical phases) in the cerebral cortex. The following adaptation of hypoxia "trained" rats to new natural conditions of Alpine altitude (3200 m) proceeded favourably, without disturbance of differentiation inhibition and without phasic states. Rats without preliminary training to altitude chamber hypoxia, in mountains (3200 m) were subjected to moderate tension resulting in protective inhibition, partial amnesia and transient disturbance of differentiation inhibition.     

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.     

Neurotic disorders in monkeys during increased stress on analytic-synthetic processes

It has been shown in experiments on three rhesus monkeys that an enhanced stress of analytical-synthetic processes consisting in learning to differentiate a sequence of tones, leads to neurotic disorders of monkeys higher nervous activity. They are manifested individually and consist in disturbances of general behaviour, in forced movements, breakdowns of differentiations, the missing of positive responses in the previously elaborated conditioned stereotype and in typical dynamics of the formation of positive and negative conditioned reflexes to a chain of tones (2000 + + 4000 c/s; 2000 + 2000 c/s; 4000 + 2000 c/s; 4000 + 4000 c/s).     

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.     

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".     

Disruption of situational reflexes in experimental neurosis brought on by changes in the space factor

Motor activity elaborated in dogs and consisting of delayed choice reactions in a stochastic system (methods developed by Beritashvili, Hunter) and of rhythmic stereotypes of motor situational reflexes (Kupalov method) was not impaired by changes in spatial parameters of unconditioned stimulus in the stochastic system of choosing one feeding trough out of five. The changes in the spatial arrangement of conditioned and unconditioned stimuli in stable stereotypes led to changes in the behaviour of the animals and experimental neurosis with long-lasting disturbances in the situational conditioned reflexes and the short-term memory.     

Tonic component of the conditioned reflex process and its functional role

Tonic background activity of 266 neurones in the hippocampus and different neocortical areas was studied in freely moving rabbits in the process of defensive and food instrumental conditioned performance and during switching-over of instrumental and classical food and defensive reflexes. Associations of CS and reinforcement evoke background activity changes in most of recorded cortical neurones preceding the development of other conditioned manifestations. Conditioned reflex was performed only after reaching the background firing rate of almost every examined neurone optimal for its realization. The performance of different conditioned reflexes was associated with different background activity levels of cortical neurones. The above mentioned data form the experimental basis for the identification of the tonic component in conditioned process which reflects tonic character of temporary connection formation and function.     

The role of the pallidum in regulating cortical activity]

Bilateral ablation of the pallidum halves the duration of extinction of conditioned motor food reflexes and contributes to 30 to 50% extinction of the electro-defensive reflexes. Pallidum functional depression by potassium chloride or novacaine leads to a temporary total depression of conditioned motor food reflexes. Depending on the frequency of pallidum stimulation, synchronization or desynchronization of the cortical bioelectrical activity is observed. Ablation of the pallidum in anaesthetized cats results in a high amplitude and low-frequency cortical activity. Injection of large doses of potassium chloride into the pallidum results in a forced running forward which lasts 30 to 40 min. The pallidum is considreed as a structure controlling the cortex activity, which takes part in the mechanisms of sensory information processes in the course of adaptive behaviour.     

The possibility of elaborating conditioned reflexes on the basis of emotional reactions evoked by electrical stimulation of limbic structures]

In experiments on cats with chronically implanted electrodes a conditioned stimulus (200 c/s tone) was paired with electrical stimulation of limbic structures, which produces pronounced emotional behaviour: defensive, aggressive, alimentary or drinking reactions. It has been shown that conditioned reactions are not elaborated on the basis of electrical stimulation, which activates alimentary, drinking or aggressive behaviour. Neither is a conditioned reflex formed to the combination of sound and stimulation of the brain structures, which inhibits alimentary behaviour. A distinct conditioned fear reaction sets in as a result of combination of sound and electrical stimulation of brain structures, which evoke defensive behaviour. The data obtained are analyzed in the light of the significance of natural requirements of the organism for the elaboration of conditioned reflexes.     

Sound reception in marine mammals depending on sound parameters and conduction pathways

The interaction of complex sounds with the body tissues of Black Sea dolphin (Tursiops truncatus) was studied by the method of instrumental conditioned reflexes with food reinforcement. The thresholds of detecting underwater acoustic signals of different frequencies for dolphin and northern fur seal (Callorhinus ursinus) were measured as a function of pulse duration under conditions of full and partial (head above water) submergence of animals into water. It was found that sound conduction through dolphin tissues was more effective than that in a northern fur seal in a wide frequency range. Presumably, the process of sound propagation in dolphin is accompanied by changes in the amplitude-frequency structure of broad-band sounds. The temporal summation in dolphin hearing was observed at all frequencies under conditions of full and partial submergence, whereas in northern fur seal it was nearly absent at a frequency of 5 kHz under the conditions of head lifting above water.     

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.     

A comparative neuropsychological approach to the analysis of hippocampal functions

Formation of conditioned switching-over of heterogeneous instrumental reflexes in dogs was more successful than in rats, which testifies about significant differences of the functional organization of analytical-synthetic brain activity in rodents and predatory animals. Experiments with lesion of the hippocampus (in rats) and the records of its electrical activity (in dogs) allowed to conclude that the hippocampus of the both types of animals belongs to the system of structures, participating in the formation of conditioned switching-over. The data, obtained in the process of elaboration of the switching-over and probabilistic reinforcement of the alimentary conditioned stimulus, testify that providing for the reactions to signals with low probability of reinforcement in different species of animals is one of the functions of the hippocampus.