Role of intralaminar thalamic nuclei in strengthening inhibition induced by stimulation of the caudate nucleus

The influence of preliminary subthreshold activation of thalamic intralaminary nuclei on achievement of instrumental reflex and on inhibitory effects, caused by stimulation of the caudate nucleus head, were studied in chronic experiments on 5 dogs with a model of instrumental defensive conditioned reflexes, providing for maintainance of a given posture. It was shown that the preceding high-frequency electrostimulation of intralaminary nuclei activates motor components of the instrumental response (shortened latency, EMG and mechanogram of the response, increased amplitude of instrumental response) and significantly lowers threshold strength of the current, necessary for obtaining "caudate pause". The greatest increase in inhibitory influences, caused by stimulation of the caudate nucleus head, was observed when thalamic stimulation preceded stimulations, localized in the dorsolateral segment of the caudate nucleus head. The obtained data are discussed in aspect of Buchwald et al. hypothesis on the existence of "caudate loop".     

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.     

The vegetative component of a conditioned reflex]

Defensive and alimentary conditioned reflexes were studied on normal cats and cats with neural isolation of the neocortex. The cardiac component of the conditioned reflexes is elaborated in decorticated animals at a similar rate and is as steady as in normal cats. Motor conditioned reactions in decorticated cats are difficult to elaborate. Both in normal and decorticated cats unconditioned pain stimulation evokes tachycardia, while a conditioned signal, paired with this unconditioned stimulus, produces bradicardia. Ban assumption has been made that the primary conditioned reaction consists in the appearance of a certain emotional state which changes the cardiac rhythm in a typical way.     

Motor alimentary conditioned reflexes and delayed reactions in dogs following destruction of the caudate nucleus]

The characteristics of formation and achievement of motor conditioned reactions were studied in seventeen dogs before and after the lesion of head or body of the caudate nucleus. It has been shown that the degree of higher nervous activity disturbances depends on the localization of injuries in the nucleus, and on the complexity of the tests used. The caudate nucleus exerts a modulating effect on the parameters of motor conditioned reflexes and takes a direct part in the structure of delayed reflexes. Lesion of the caudate nucleus body is attended with more serious disturbances of higher nervous activity than lesion of its head.     

Inhibitory effect of the amygdala on alimentary-conditioned reflexes in the dog

In experiments on dogs, changes of conditioned alimentary reflexes were studied under stimulation (50-100 mcA, 1, 40, 100 imp/s, 0.5 ms) of the basolateral area of amygdala before and after electrolytic ablation (5 mA, 60 s) of the head of the caudate nucleus. Stimulation of the amygdala, which elicited inhibition of conditioned secretory reflexes and prolongation of latencies in intact dogs leads to opposite - activating - influences after destruction of the caudate nucleus head. A conclusion is made on the participation of neurophysiological and neurochemical mechanisms of the caudate nucleus in transmission of inhibitory effects of the amygdala on alimentary conditioned reflexes.     

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.     

Effect of electric stimulation of different portions of the amygdala on the formation of alimentary conditioned reflexes in the dog

In dogs with previous learning experience, a low-frequency (1 imp/s), weak (20-50 mcA) amygdala stimulation produces dissimilar influences on formation rate and value of conditioned alimentary reflexes in dependence on stimulation localization. At stimulation of amygdala basolateral part, the reflexes formation proceeds 2-3 times slower than normally. Their value is significantly reduced, the triggering of the secretory reaction is delayed. Dorsomedial part stimulation accelerates the process of conditioning, intensifies the secretion but delays the start of secretory reaction. The elaboration of differentiation inhibition is facilitated by basolateral part stimulation and is impeded by dorsomedial amygdala activation.     

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.     

Electrophysiologic and behavioral changes after destruction of the basolateral nucleus of the amygdala in the rat

In experiments on rats the influence was studied of unilateral destruction of the basolateral amygdala on heterogeneous conditioned reflexes and focal potentials (FPs) of the dorso-medial thalamic nucleus, elicited by stimulation of the peri-amygdalar cortex. It is shown that reduction during 3-4 postsurgery weeks of the number of goal-directed reactions and increase of their latencies are accompanied by suppression of the evoked activity, testifying to disturbance of functional interaction between the structures of limbic and thalamic levels. Changes of FPs parameters, reflecting the establishment of new functional interrelations between the centres of the lesioned system excelled in duration compensatory restoration of properties of conditioned reflexes.     

Conditioned reflex reproduction of evoked potentials during reinforcement by electrostimulation of the hypothalamus]

Conditioned evoked potentials to stimulation of a limb, a light flash or a short tone were studied on rats in acute experiments with nembutal anaesthesia and in chronic experiments. Electrical stimulation of the hypothalamus was used as reinforcement. Following the pairings of conditioned and hypothalamic stimulation, the appearance of conditioned EP was recorded, as manifest in the period of the previously applied unconditioned signal after its omission. Their emergence was facilitated after high-frequency stimulation of the hypothalamus or administration of galantamine, an anticholinesteraze drug. Coagulation of the hypothalamus did not prevent any manifestation of conditioned evoked potentials.     

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.     

Interaction between depressor and pressor reflexes arising from the aorta in dogs

In 18 dogs anesthetized with morphine-chloralose the interaction between the aortic nerve (AN) pressor and depressor reflexes was studied. Low-intensity, high-frequency electrical stimulation of the AN causes large decreases in heart rate and systemic pressure characteristic of baroreflex responses. High-intensity, low-frequency stimulation of the AN causes modest increases in heart rate and systemic pressure similar to the responses observed to intraaortic nicotine. Simultaneous electrical stimulation of these antagonistic reflexes results in much smaller (P less than 0.001) reductions in heart rate and systemic pressure that can be explained on the basis of simple addition of the individual responses. Similarly the AN depressor reflexes are suppressed during intraaortic infusions of nicotine (40 micrograms/min). The results suggest that the inhibitory effects of the AN baroreflexes are suppressed by the aortic chemoreflexes . This interaction occurs in the CNS rather than at the level of the heart or vascular smooth muscle.     

The role of the lateral and medial hypothalamus in reproducing food and escape instrumental reactions]

In experiments on two dogs it has been found that testing electric stimulation of the lateral hypothalamus reproduces instrumental movement in case of elaborated alimentary instrumental conditioned reflexes (CR) to a tone and does not reproduce it at elaborated acid-defensive instrumental CR. Under testing electric stimulation of the medial hypothalamus the instrumental movement practically is not reproduced neither in alimentary nor in defensive situation. Appearance of instrumental movement at LH electro-stimulation in alimentary situation is connected with activation of backward conditioned connection from motivation "alimentary" LH centre to representation of this movement in the motor cortex.     

Generalization of an instrumental defensive conditioned reflex elaborated to electric stimulation of the hypothalamus in dogs

Generalization of defensive conditioned reflexes elaborated to electrical stimulation of the lateral hypothalamus (LH) was studied in four dogs with electrodes implanted in various structures of the limbic system. Electrocutaneous stimulation was switched off when the dog lifted the foreleg to a definite level. Generalization of the conditioned reflex was manifested in different degrees when testing different formations of the limbic system, or testing one and the same structure, but at different stages of conditioning. Two types of generalization were found: the first one--a well pronounced motor reaction, by its latency, level and duration of lifting the foreleg similar to movements appearing in response to the conditioned stimulus; and the second one--low amplitude or short-term movements differing from conditioned ones. The first type of generalization was observed in response to stimulation of LH, contralateral to the point of signal stimulation, of the ventromedial hypothalamic nucleus, the mammillary bodies and the basal, lateral and to a lesser degree, the central nucleus of the amygdaloid complex; the second type--in response to stimulation of the ventral hippocampus, the medial and lateral septum nuclei.     

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.     

The role of the medial geniculate bodies in analyzing sounds]

Elaboration and manifestation of motor alimentary conditioned reflexes were studied after a lesion of medial geniculate bodies (MGB) in adult cats. No substantial disturbance were recorded of the capacity for pitch analysis. In the case of more extensive MGB lesions, there were some difficulties in the elaboration of tone differentiations but not in their achievement. Additional temporo-parietal cortical ablation drastically reduced the animals' capacity for anlysing sound frequencies and more profoundly disturbed their capacity for inhibitory reactions.     

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.     

Transformation of conditioned responses of the tonic type

A reversal of signal meaning of stimuli signalling experimental situation was performed on dogs with conditioned defensive motor reflexes to clicks: clicks and electro-cutaneous stimulation were stopped, and the animals were started to be given food in the same experimental situation. Reversal of conditioned defensive tonic stimuli to alimentary ones led to a complex of behavioral and electrographic reactions. Isolated click tests showed that signals of experimental situation considerably affect a fully-elaborated defensive motor reflex. At the same time the preparatory role of certain situational elements may be specialized with regard to the components of the conditioned reflex, in particular to the local flexor reaction.     

Physiologic characteristics of systems of conditioned connections during urgent formation of a program of action]

Examination of preschool age children has shown that instantaneous formation of a system is possible only on the basis of integration of conditioned reflexes, which are differentiated by the nature or strength of reinforcement. Integration of previously elaborated conditioned reflexes is more successful if the initial systems of conditioned connections were in a state of conditioned tonic excitation of an optimal strength. Uniformly strong or uniformly weak stability of all reflexes in the system renders it inert, hence makes it a poor basis for instantaneous formation of a new system.     

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.