Significance of the relative position in space of associated stimuli for formation and performance of motor conditioned reflexes

In experiments on cats the rate of formation of conditioned reflexes to sound (running to the feeding trough) depended on the spatial interrelations of the paired stimuli: the closer the source of the conditioned signal to the feeding trough, the sooner the formation of the conditioned reflex. It has been assumed that during formation of a conditioned reflex the closing of connection between conditioned and unconditioned stimuli is also attended with the closing of connection between the spatial parameters of the paired stimuli. Experiments with inactivation through cold of the temporal area (cortical representation of the vestibular system) of one hemisphere have shown that such a connection is formed in central parts of the vestibular analyser. When the conditioned reflex is elaborated to one feeding trough, the connection is duplicated by both hemispheres; in reflexes to two feeding troughs (i.e. spatial choice) such connections are lateralized in each hemisphere.     

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.     

Relation between the level of conditioned reflex activity and concentration of serotonin in the blood of normal and neurotic dogs

Higher nervous activity of dogs was studied using Pavlovian method of alimentary salivary conditioned reflexes. Serotonin (5- hydroxytryptamine) content in the venous blood was defined by means of fluorescent spectrofluorimeter using orthophthalic dialdehyde. A high level of conditioned and unconditioned reflexes was accompanied by a low serotonin blood content, while a low level of reflexes was accompanied by a high serotonin content. Negative correlation is supposed to exist between the degree of activation of nervous processes and blood serotonin content. Blood serotonin content increased in dogs in experimental neurosis with expressed reduction of the level on alimentary conditioned and unconditioned reflexes.     

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".     

Features of the formation of conditioned secretory reflexes in response to the stimulation of different areas of the caudate nucleus of the dog

Properties were studied of elaboration of conditioned alimentary secretory reflexes to threshold electric stimulation of the caudate nucleus in dogs. The rate of formation of such a reflex and its magnitude were considerably lower than in reflexes elaborated to distant stimuli. Latencies were longer; dynamics of conditioned and unconditioned secretory reactions was similar to that of reflexes to a weak stimulus--light. The motor component of the conditioned alimentary reaction, as a rule, was absent. Characteristics were revealed of formation of alimentary reflexes to stimulation of the dorsal and ventral parts of the head and body of the caudate nucleus.     

Conditioned reflexes before and after lesioning the substantia innominata in the cat

It has been shown that bilateral lesion of the substantia innominata has caused temporary depression of unconditioned and conditioned alimentary reflexes, disturbances of the elaboration and performance of conditioned reflexes. It is suggested that the latter produced by the operative memory disturbances.     

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.     

Effect of activating the putamen on alimentary conditioned reflexes in dogs

The influence was studied of the near-threshold high- and low-frequency stimulation of the putamen on achievement of alimentary conditioned reflexes in dogs. Both low- and high-frequency activation of the examined structure significantly changes the parameters of intermediate secretion, of the salivary conditioned reflexes, without influencing unconditioned ones. Unilateral stimulation disturbs the achievement of positive and inhibitory conditioned reflexes to a greater extent than bilateral one. Difference between the effects of low- and high-frequency stimulation has been revealed.     

Conditioned alimentary motor reflexes in cats following destruction of the scorpula]

Bilateral ablation of the putamen results in temporary disappearance of natural as well as artificial conditioned motor reflexes in cats. Further training is necessary for their reestablishment. Restored conditioned reflexes appear with a long latency. After partial bilateral ablation of the putamen, animals that have been trained not only to press a pedal, but also to make a choice between a right or left feeding tray, lose the latter ability. After the reestablishment of conditioned reflexes, the animal makes a correct choice of side only in 68--70% of cases. After total bilateral ablation of the putamen the animals lose artificial conditioned reflexes. The results suggest that the putamen has a close relation to the mechanisms of learning and memory.     

Vector representation of associative learning

I. P. Pavlov [12] has shown that conditioned reflexes are selective both with respect to conditioned stimuli and to conditioned reflexes elicited by those conditioned stimuli. At the neuronal level selective aspects of conditioned stimuli are based on detectors selectively tuned to respective stimuli. The selective aspects of conditioned reflexes are due to command neurons representing specific unconditioned reflexes. It can be assumed that conditioned reflexes result from association between selective detectors and specific command neurons. The detectors activated by a conditioned stimulus constitute a combination of excitations--a detector excitation vector. The detector excitation vector acts on a command neuron via a set of plastic synapses--a synaptic weight vector. Plastic synapses are modified in the process of learning making command neuron selectively tuned to a specific conditioned stimulus. The selective tuning of a particular command neuron to a specific excitation vector referred to a conditioned stimulus is a basis of associative learning. The probabilities of conditioned reflexes elicited by conditioned and differential stimuli implicitly contain information concerning excitation vectors that encode respective stimuli. Contribution of the vector code to associative learning was explored combining differential color conditioning with intracellular recording from color-coding neurons. It was shown that colors in carps and monkeys are represented on a hypersphere in the four-dimensional space similar to human color space. The basis of the color space is constituted by red-green, blue-yellow, brightness and darkness neurons.     

Reproduction of the electrophysiologic correlates of a conditioned defense reflex following electroconvulsive activity]

Acute and chronic experiments on cats have shown the possibility of obtaining in one session conditioned EEG-responses and vegetative changes as well as conditioned motor reactions time-locked to the moment of the unconditioned pain stimulation omitted during testing. Electro-convulsive stimulation directly after elaboration prevents the appearance of conditioned motor reactions, while conditioned EEG and vegetative changes are retained after electric seizures. A conclusion has been drawn that the emotional component of conditioned reactions is not subjected to the amnestic effect of electro-convulsive action.     

Inhibitory effects of external stimulation on neuronal spike response in the cat association cortex during conditioning

The effects of external stimulus presentation on neuronal spike response induced in area 7 by conditioned and unconditioned stimuli were investigated in waking cats. Pointing reaction arising in response to the action of external stimuli led to the phasing out of spike response to unconditioned cutaneous stimulation, as well as depression or complete suppression of response in area 7 neurons to conditioned stimulus presentation, despite occurrence of conditioned reflex motion. External stimuli no longer exerted an inhibitory effect on the neuronal spike response induced by protracted use of conditioned and unconditioned stimuli. The fact that the conditioned lever-pressing reflex could occur either following or without neuronal spike response in area 7 would raise the question of whether neurons belonging to this area always participate directly in the triggering and occurrence of conditioned reflex motion.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziogiya, Vol. 22, No. 2, pp. 147–155, March–April, 1990.     

The influence of electrocoagulation of the septum and section of the entorhinal cortex on general behaviour and memory in cats

In chronic experiments on cats it was shown that the lesion of the medial part of the septum does not result in the "septal syndrome"; the ratio of the different sleep-wakefulness cycle phases remains within the baseline values; the acquisition, retrieval and extinction of instrumental alimentary conditioned reflexes proceed normally; the delayed conditioned reflexes are impaired. Massive septal lesion, including its lateral part, leads to the development of the "septal syndrome"; there are changes in the structure and percentage of the different sleep-wakefulness cycle phases; the acquisition and extinction of instrumental alimentary reflexes with the sound discrimination are markedly retarded; the performance of delayed conditioned reflexes is completely destroyed. The section of the entorhinal cortex produces an increase in the number of repeated errors and perseverative movements during performance of instrumental alimentary reflexes, the deceleration of the acquisition and extinction of instrumental alimentary reflexes, complete disturbance of the delayed conditioned reflexes and does not affect the sleep-wakefulness cycle. The problems of the role of the hippocampus and its main inputs in the regulation of the short-term operative memory in the "pure form" as well as the significance of the descending influence of the hippocampus on the regulation of general animal behavior were also discussed.     

The effect of mixtures of caffeine, glucose and ascorbic acid on conditioned reflex activity in dogs during the postanesthetic period]

It was shown in experiments on four dogs by the conditioned method that the period of recovery of conditioned activity after one hour ether anaesthesia tested 7 to 7.5 days. Premedication with a mixture of caffeine, glucose and ascorbic acid reduced the recovery time to four or five days. After the administration of the complex in the postanaesthetic period (in 2.5 hours after anaesthesia) the stabilizing effect was still more pronounced. Thus, the recovery time of unconditioned and conditioned reflexes and differentiation was reduced to three or four days.     

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.     

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 neuronal responses of area 7 of cat cerebral cortex to unconditioned and conditioned stimulation under the effects of external stimuli

The effects of external stimuli (ES) and feeding motivations on area-7 neuronal impulse responses (IR) elicited by unconditioned and conditioned stimulation in alert cats were investigated. In untrained cats, preliminary (by 1 sec) action of ES resulted in the disappearance of impulse responses to electrical stimulation of the forepaw (EFS) in the first (no more than five-eight) executions; the responses gradually reappeared and in subsequent executions reestablished their former magnitude. The emergence of feeding motivation excitation in the cats, elicited by food presentation, also promoted disappearance of the initial responses to EFS; these were reestablished only after the cats were quieted. In conditioned-reflex cats, the action of the ES, which had been suppressed after onset of the reflex, as well as lowering of initial feeding motivation level (caused by natural satiation of the cats or by change in quality of the reinforcing food item), led to increase in latent periods (by 50–250 msec) and prolongation of neuronal responses to the conditioning stimulus (CS); but these parameters were unchanged during performance of trained movements. Neuronal response lag time to the action of the CS was defined by ES delivery time and by level of feeding excitation in the cats. The reason for the variation of neuronal responses to unconditioned and conditioned stimulation in the parietal cortex is assumed to be associated with variation in exteroreceptive attention level of cats under the effects of stimuli and altered metabolism.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 23, No. 5, pp. 564–574, September–October, 1991.     

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.     

Experimental neuroses in dogs caused by an increase in the intensity of (light) stimuli conditions

The influence of enhanced intensity of general illumination on conditioned reactions of a different degree of complexity was studied on dogs by the classical method of secretory conditioned reflexes. In most of the dogs, increase of general illumination from 30 to 300 luxes leads to more intensive secretory conditioned and unconditioned reactions, a smaller number of errors in differentiation, and stabilization of parameters of conditioned activity, which testifies to the improved general functional state of the brain. In two animals in the same conditions of the experiment, an experimental neurosis set it, not connected with the complexity of the elaborated activity. It is assumed that the neurotizing agent was the superfluous situational (photic) stimulation which presented excessive requirements to the mechanisms regulating the general functional state of the brain.     

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.