Integrative properties of parietal cortex neurons of rabbits during the switching of heterogeneous classical conditioned reflexes

The impulsive activity of parietal cortex neurons of rabbits during the elaboration and specialisation of switching-over of alimentary and defensive conditioned reflexes has been studied. There are changes of parietal cortex neurons activity correlated with the stages of switching-over elaboration. The significant differences of the background neurons activity and neurons response to the conditional signal during alimentary and defensive conditioned reflexes have been established. The dominant motivation influences the neurons activity by modulation of background and conditioned activity of neurons. The integrative properties of neurons investigated in switching-over situation are revealed in their ability to summarize the excitation coming to the same sensory input with excitation from different motivational and reinforcement systems.     

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.     

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 typological characteristics of higher nervous activity in dogs and the maxima of the cross-correlation function between the electrical activities of the frontal cortex and the brain limbic systems]

Electrical activity of the frontal cortex, dorsal hippocampus, basolateral amygdala and lateral hypothalamus was recorded in eight dogs with chronically implanted electrodes. Mean values of the maxima of crosscorrelation function (MCCF) between electrical potentials in the theta, alpha and beta-2 ranges were used as a basis for assessment of conditions for interaction between these structures. Typological features of the higher nervous activity were assessed by the animal performance under conditions of free choice of the reinforcement mode of a conditioned stimulus: either high probable but of low alimentary quality or with low probability but more valuable. The mean MCCF values in the theta range were higher than in the other ranges. The brain structure which had the high MCCF in the theta-range, at least, with two of the structures under study was considered as "dominant". It was shown that hippocampus was the dominant structure for melancholic dogs, the frontal cortex was in phlegmatics. The hypothalamus was shown to be the "dominant structure" in both sanguine and choleric animals, but, for the most part, its activity was correlated with different structures. Thus, conditions for interaction between the frontal cortex, hippocampus, amygdala and hypothalamus seem to be an important factor, which determines typological features of the higher nervous activity of dogs.     

Conditioned reflex and electrical activities of the hippocampus during the random use of heterogeneous triggering stimuli

In dogs with electrodes chronically implanted in the hippocampus, the conditioned activity, heart rate frequency and spectral characteristics of the hippocampal theta-rhythm were studied in conditions of irregular presentation after a preparatory stimulus now of alimentary and now of defensive triggering conditioned stimuli. It is shown that uncertainty of alimentary or defensive reinforcement is a stronger emotiogenic factor than the action of the triggering defensive conditioned stimulus. Changes in electrical hippocampal activity and autonomous activity depend, along with other factors, on forecasted volume of the forthcoming motor activity. The speed of instrumental conditioned reflexes formation correlates with the hippocampal theta-rhythm frequency, typical for the given animal.     

The dependence of hippocampal electrical activity on the probability of the reinforcement of a conditioned food stimulus

In three dogs the dynamics was studied of changes in the number of instrumental motor reactions, the heart rate and the hippocampus theta rhythm was studied at the change of constant reinforcement to various regimes of probable reinforcement of alimentary conditioned stimulus. At 70 and 50% levels of the reinforcement the instrumental reflex appeared in response to all presentations of the signal stimulus. The greatest decrease in the number of reactions took place at 5% probability. The heart rate both in the intersignal period and at getting of empty feeder depended on the used probability of reinforcement and individual characteristics of animals. The frequency of the theta rhythm in all dogs was the highest at the 30% level and the lowest values were found at the 5% level of reinforcement. The obtained facts testify once more to complex genesis of emotional excitation, the value of which depends both on informational (uncertainty) and motivational (significance of reinforcement) factors.     

Ontogenetic characteristics of the acquisition and extinction of a taste aversion in dogs following damage to the dorsal area of the hippocampus

In 2-9 months dogs the influence was studied of ablation of the hippocampus dorsal area on formation and preservation of conditioned taste aversion (CTA), elaborated by combination of 30% sucrose solution with i.p. injection of 0.28 M LiCl solution. Lesion of the hippocampus dorsal area does not prevent acquisition after the first pairing of conditioned taste aversion in puppies and adult dogs. Heterogeneous influences are observed after hippocampus lesions on the process of CTA extinction in animals of different ages. Acquaintance with conditioned stimulus before CTA acquisition accelerates the process of its extinction in hippocampectomized indiviuals, but less than in animals with an intact hippocampus.     

Patterns in the transition from normal to neurosis in dogs with different typological traits in food and defensive situations

At comparison of the formation of neurosis in dogs with different typological characteristics in conditions with alimentary and pain reinforcement, a resemblance was revealed of transition from norm to the initial stage of pathology. In strong animals a development was seen of compensatory tachycardia at circular changes of conditioned activity. Weak type dogs are characterized by a lowering or full disappearance of reflexes in combination with bradycardia. The method of differentiation, allowing in situations with alimentary reinforcement to classify the tested animals according to the type of higher nervous activity and differentiate the states of "norm" and "initial stage of neurosis", proved to be effective for the analogous aims also in studies with defensive reinforcement.     

The individual organization of frontal-hippocampal networks during realization of different behavioral tasks

Four cats were subjected to appetitive instrumental conditioning with light as a conditioned stimulus by the method of "active choice" of the reinforcement quality: short-delay conditioned bar-press responses were followed by bread-meat mixture and the delayed responses--by meat. The animals differed in behavior strategy: four animals preferred bar-pressing with long delay (so called "self-control" group); two animal preferred bar-pressing with short-delay (so called "impulsive" group). Then all the animals were learned to short-delay (1 s) instrumental conditioned reflex to light (CS+) reinforced by meat. The multiunit activity in the frontal cortex and the hippocampus (CA3) was recorded through chronically implanted nichrome-wire semimicroelectrodes. The interactions among the neighboring neurons in the frontal cortex and hippocampus (within the local neuronal networks) and between the neurons of the frontal cortex and hippocampus (distributed neuronal networks of frontal-hippocampal and hippocampal-frontal directions) were evaluated by means of statistical crosscorrelation analysis of the spike trains. Crosscorrelation interneuronal connections in the delay range 0-100 ms were explored. It was shown that the functional organization of the frontal and hippocampal neuronal networks differed in choice behavior and was similar during realization of short-delayed conditioned reflex. We suggest that the local and distributed neural networks of the frontal cortex and hippocampus take part in the realization of cognitive behavior, in particularly in the processes of the decision making.     

Conditioned reflex switching over in unrestrained dogs

Conditioned switching-over was studied in unrestrained animals by means of instrumental alimentary conditioned reflexes of pressing the pedal. A complex of auditory and spatial factors (start-ground) served as conditioned alimentary signal. Backward connections were stable and were manifested in the teturning of the animal to the start-ground after feeding. It has been confirmed that the observed working readiness is a conditioned reflex of a tonic type.     

The effects of intracerebral injections of kynurenine on conditioned reflex activity and on the serotonin level of the blood in dogs]

Higher nervous activity of dogs was studied by classical Pavlovian method of alimentary salivary conditioned reflexes and serotonin content in blood was measured. Kynurenine sulfate in a dose of 300 + 300 mkg (bilaterally) was injected in the dorsal hippocampus region through chemiotrodes. In most cases the injection of the substance led to changes towards excitation (increase of conditioned and unconditioned alimentary salivation in the day of injection and the next day) and also to a rise of serotonin content in animals blood.     

Effect of destruction of the hippocampus and septum on learning in the rat in "negative transfer" situations

Intact rats and rats with lesions of the hippocampus and septum were trained to repeated switching-over of the habit of choosing the reinforced side of the maze as well as to alternation of reinforced side choice. Repeated switchings-over of the habit impedes the subsequent elaboration of the choice alternation ("negative transfer"). In operated animals, the interferating effect of the previous experience is more expressed. The observed deficit of training is due to disturbance in the apparatus limiting the process of search and retrieval of adequate information from the memory.     

Increased regularity of activity of cortical neurons in learning due to disinhibitory effect of reinforcement

This paper reviews the author's studies on neurophysiologic mechanisms of conditioned reflex learning. Electroencephalograms, evoked potentials, activity of neocortical and hippocampal neurons and the rabbits' behavior in the course of elaboration of defensive and inhibitory conditioned reflexes to light flashes have been recorded. Electric shock (ECS) applied to the paw served as reinforcement. The study demonstrated three types of reinforcement effect on the activity of cortical neurons: activating, disinhibitory, and inhibitory. EEG activation due to reinforcement is accompanied by a change in phasic cortical neuronal activity from chaotic or irregular, typical of rest or inhibition, to regular tonic discharges (in neocortex and hippocampus) and group discharges in the stress rhythm, 5-7 Hz in the hippocampus. Following a number of conditioning trials, the effect of reinforcement is simulated by the effect of a conditioned stimulus. With EEG activation and increased regularity in impulses, facilitation of motor reactions is observed.     

Interaction between neurons of the frontal cortex and hippocampus during the realization of choice of food reinforcement quality in cats

Six cats were subjected to the procedure of appetitive instrumental conditioning (with light as a conditioned stimuls) by the method of the "active choice" of reinforcement quality. Short-delay conditioned bar-press responses were rewarded with bread-meat mixture, and the delayed responses were reinforced by meat. The animals differed in behavior strategy: four animals preferred the bar-pressing with a long delay (the so-called "self-control" group), and two cats preferred the bar-pressing with a short delay (the so-called "impulsive" group). Multiunit activity in the frontal cortex and hippocampus (CA3) was recorded via chronically implanted nichrome wire semimicroelectrodes. An interaction between the neighboring neurons in the frontal cortex and hippocampus (within local neural networks) and between the neurons of the frontal cortex and hippocampus (distributed neural networks in frontal-hippocampal and hippocampal-frontal directions) was evaluated by means of statistical crosscorrelation analysis of spike trains. Crosscorrelations between neuronal spike trains in the delay range of 0-100 ms were explored. It was shown that the number of crosscorrelations between the neuronal discharges both in the local and distributed networks was significantly higher in the "self-control" cats. It was suggested that the local and distributed neural networks of the frontal cortex and hippocampus are involved in the system of brain structures which determine the behavioral strategy of animals in the "self-control" group.     

Effects of Fat-Enriched Diet on the Content of Sphingolipids in the Brain and on Cognitive Functions in Old Rats

It was demonstrated that levels of free fatty acids and radioactively labeled ([¹⁴C]) ceramide in the liver, muscles, and brain of rats fed a saturated fat-enriched diet for 36 weeks (from 15 to 24 months) increased, as compared with the corresponding levels in control animals of the same age (24 months old). A high-saturated fat diet resulted in a rise in the content of sphingolipids synthesized de novo in the neocortex and hippocampus of 24-month-old rats. High levels of ceramide and sphingosine in the neocortex and hippocampus induced in these rats by the above diet correlated with considerable worsening of the conditioned reflex activity of animals (reflex of active avoidance in the shuttle chamber).     

The long-term effect of acute hypoxic hypoxia on shuttlebox avoidance learning in rats

Long-term influence was studied of the acute hypoxic hypoxia seance on rats behaviour in situation of elaboration of the conditioned reaction of active avoidance of electric shocks in shuttle chamber. It was found that in 2.5-3 months after the hypoxia seance, the experimental animals significantly differed from the intact controls by dynamics of CR elaboration (rats which had hypoxia were ahead of the control ones) and by distribution of the conditioned reactions latencies (for experimental animals this distribution was shifted to minor values). The character of these behavioural shifts coincided with that observed in the group of rats with local unilateral hippocampus lesion. The obtained results and numerous data presented in literature on the influence of the hippocampus lesion on animals shuttle avoidance learning, allow to conclude that the seance of hypoxic hypoxia leads to the disturbance of the hippocampus function. This conclusion conforms to the data on diffusive death of the hippocampal and neocortical neurones as a result of hypoxia action.     

Characteristics of memory and the functional organization of the hippocampo-reticulo-neocortical complex of the brain in the acute postoperative period

In tests on dogs, rabbits and rats, it has been established that in acute postoperative period, the time of memorizing of conditioned signals (light, tone and metronome) is reduced, the time of realization of conditioned reactions increases, the excitability and the bloodflow of the midbrain reticular formation and Mg-AtPhase activity of pons Varolii raise. In the hippocampus the excitability and local bloodflow lower and the activity of Ca-Mg-ATPhase is enhanced. In the frontal cortex these processes do not change. Functional interrelations of the brain structures in the acute postoperative period are characterized by the weakening of the activating influence of the reticular formation on the frontal cortex and an increase of its suppressive action on the dorsal hippocampus. It is suggested, that the discovered damages in the higher nervous activity are stipulated by the changes in neurochemical organization of the brain.     

Effect of tuftsin and its analog on learning, memory and exploratory behavior in rats

On Wistar male rats the influence of tetrapeptide tuftsin and its analogue TP-1 (300 mkg/kg, i/p) on animals behaviour was studied. It is shown that administration of peptides increases the rat's exploratory activity. Activating influence of TP-1 was continued for 6 h. Daily multiple administration of tuftsin or TP-1, 15 min prior to the beginning of experiments facilitates the learning and stability of conditioned reaction with food reinforcement. Experimental animals react significantly weaker than the control ones to emotionally negative influence, produced by a sharp reduction of the amount of alimentary reinforcement control.     

The relation of the typological classification of dogs to their behavioral characteristics in variations between the probability and the value of reinforcement]

In studying of dogs behaviour in coordinates system "Requirements and probability of their satisfaction" it has been established that selection of behaviour strategy is closely connected with individual typological properties of animals. In experiments with equally-probable alimentary reinforcement a direct correlation was found between the strength parameter of the nervous system and the strategy of dogs behaviour. Animals of strong types manifested the character of reaction which consisted in determination and goal-directedness, while the animals of the weak type manifested other character, having features of uncertainty, indefinition, hesitations. In conditions of conflict between probability and value of reinforcement the dogs manifested two opposite strategies of behaviour: orientation to highly probable events (choleric and phlegmatic) and to low-probable events (sanguinic and melancholic) what is connected with individual properties of functioning and the character of interaction of four brain structures (frontal cortex, hippocampus, hypothalamus, amygdala).     

Individual typological characteristics of higher nervous performance and asymmetry of hippocampus and amygdala electrical activity in dogs

Electrical activity of the frontal cortex, dorsal hippocampus, basolateral amygdala and lateral hypothalamus of both hemispheres was recorded in nine dogs in the state of quiet wakefulness without any stimulation. Individual typological features of higher nervous activity were assessed by the animal performance under conditions of free choice of the reinforcement mode: either high probable but of low alimentary quality, or with low probability but more valuable. Mean values of the maxima of crosscorrelation function between electrical activity of the investigated structures of two hemispheres were used as a basis for assessment of conditions for interaction between left and right formations. For the hippocampus and amygdala, in some dogs these conditions were the best in the theta and beta 2 ranges, in other animals--in the theta and alpha bands. In phlegmatic dogs, spectral densities in the theta range were higher in the left hippocampus than in the symmetrical structure, in sanguine animals spectral densities in the theta and beta 2 ranges in the hippocampus and amygdala were higher in the right hemisphere than in the left one. Thus, the hemispheric asymmetry of electrical activity of the limbic formations seems to be an important factor, which determines the individual typological features of the higher nervous activity in dogs.