Neuronal response in the cat parietal association cortex to conditioned and non-conditioned acoustic stimuli

Neuronal response in the cat association cortex (area 5) to conditioned and non-conditioned acoustic stimulation was investigated. Numbers of neurons responding to a conditioned acoustic stimulus according to the traditional reflex pattern were twice as high. Numbers of inhibitory neuronal responses to the stimulus increased when instrumental reflex occurred. Neurons were found which only reacted to a conditioned acoustic stimulus in the absence of conditioned reflex movement occurring with instrumental food reflex. Although findings do not exclude the possibility of this cortical area contributing to the analysis of sensory signals and evaluation of their biological significance, it might be supposed that its main functional property lies in its involvement in the process of initiating behavioral response to a conditioned response.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 20, No. 5, pp. 637–645, September–October, 1988.     

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.     

Neuronal response in the cat parietal association cerebral cortex (area 5) during performance of conditioned reflex motion

Responses in 160 neurons of the cat parietal cortex were investigated during the performance of instrumental food reflex (lever pressing) during experiments involving presentation of a conditioned acoustic stimulus. Discharge rate changed in 49% of neurons during the period preceding the conditioned reflex movement. Three basic types of cell with an excitatory response pattern were discovered apart from a small group showing suppression of activity, each differently involved in the process of conditioned reflex movement performance. Excitation arose in neurons of the first type 200±52.9 msec (average) before the onset of the conditioned reflex movement, reaching its peak discharge rate as the animal placed its paw on the lever. The former parameter was 605±54.2 msec for the second type of neuron, with firing rate peaking between the start of electromyographic response and the completion of lever pressing. The same parameter measured 1,000–2,000 msec in the third type and activation took the form of a diffuse increase in discharge rate without a clear-cut peak occurring during performance of the instrumental reflex. Findings would suggest the involvement of the parietal cortex neuronal system in the triggering as well as the follow-through of conditioned reflex motion.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 19, No. 2, pp. 223–231, March–April, 1987.     

The conditioned avoidance reflex and intersignal movements: the correlation of the cardiac and motor components]

In five dogs correlation was studied of heart rate (HR) and motor component of conditioned avoidance reflex (CAR), elaborated by Petropavlovski? method. Stable CAR was expressed in lifting and long (not less than 5-10 sec) holding of the paw on definite height (5-10 cm) for the avoidance of painful electrocutaneous paw stimulation in response to conditioned acoustic stimulus. The level of defensive excitation, evaluated by heart rate change was maximum before the beginning of the conditioned motor reaction. Immediately after lifting and placing the paw in the zone of security a sharp decrease of defensive excitation level ("drive" reduction) took place. Intertrial motor reactions of two types were revealed. The first type imitated the conditioned motor reaction, the second one the usual phasic bending of the paw. Against the background of intertrial movement of the first, operant type a decrease of defensive motivation took similarity as it occurred against the background of CAR during the performance response.     

Role of the commissural channels of the somatosensory system in the conduction of feedback signals during the control of voluntary movements

The time-course of formation of the conditioned defence reflex and electromyograms of the working forelimb were studied on cats with classical and commissural somatosensory pathways transections. It was established that exclusion of the classical somatosensory projections only reduces the rate of the skill formation that is related to the possibility of conducting feedback signals via the remaining commissural pathways. However, the commissural channel in question is not effective enough, since its isolated functioning is accompanied by an increase in the duration of instrumental movement. The combined transection of the classical and commissural somatosensory pathways excludes the possibility of formation of the conditioned reflex on the "deafferented" limb. The lack of direct visual control over the moving forelimb does not essentially affect the behavioral and electromyographic data.     

New data on the correlation of the cardiac and motor components of the instrumental conditioned defensive reflex in dogs

Correlation of heart rate (HR) and conditioned motor reaction (CMR) was studied in dogs with conditioned avoidance reflex by the V. P. Petropavlovski? method. An increase of HR was shown in response to presentation of conditioned sound stimulus reaching maximum directly before CMR beginning. Immediately after the beginning of CMR and further against the background of continuous keeping of the paw in the safety zone, CMR greatly decreases. The CMR changes testify to a heavy weakening of classical conditioned reflex of fear as a result of kinesthetic afferentation signalling about successful keeping of the paw in the safety zone.     

Role of the midbrain periaqueductal gray matter in conditioned reflexes

Unit activity in the midbrain periaqueductal gray matter (PGM) during an instrumental placing reflex, its extinction, differentiation, and conditioned inhibition, was studied in chronic experiments on cats. Spike responses 1–2 sec in duration in 69 (36.7%) of 182 neurons preceded by 400–800 msec the beginning of conditioned-reflex and voluntary intertrial movements. These advanced responses appeared 200 msec before the corresponding advance responses of motor cortical neurons. Fifty-eight neurons (30.9%) responded directly to acoustic stimulation with a latent period of 10–50 msec for 2–6 sec, 19 neurons (10.1%) generated double responses, linked with both the acoustic stimulus and subsequent conditioned-reflex movement, and 42 neurons (22.3%) did not respond to acoustic stimulation, although individual neurons of this group changed the level of their spontaneous activity in response to repeated conditioned stimulation, and this change was maintained for some tens of minutes. Extinction, differentiation, and conditioned inhibition all abolished conditioned-reflex movements, but each type of internal inhibition was accompanied by its own characteristic changes in the firing pattern of PGM neurons. Functional independence of neurons of the first and second groups was demonstrated during extinction and recovery of the conditioned-reflex. The results indicate the important role of PGM not only in the mechanism of the conditioned reflex, but also in the development of its internal inhibition.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 16, No. 3, pp. 403–419, May–June, 1984.     

Responses of motor cortex neurons of the cat to auditory stimulation and their role in the instrumental food reflex

The responses of motor cortex neurons in the cat to the presentation of a single auditory click and a series of 10 clicks presented with 1,000/sec frequency were studied under conditions of chronic experiments before and after the development of an instrumental food reflex. After reflex development a single presentation of a positive conditioned stimulus (single click) markedly influenced for 7 sec the appearance of instrumental movements. At the same time, the immediate responses of motor cortex neurons to presentation of the conditioned auditory stimulus had no impact on the appearance in the motor cortex of discharges leading to the realization of instrumental movements. Consequently, motor cortex neurons do not require activation from afferent sensory inputs for the generation of such discharges. The immediate neuronal responses to conditioned stimulation did not inhibit the realization of the instrumental reflex. It is proposed that they are associated with the realization of motor function in the unconditioned defensive response evoked by the presentation of an auditory stimulus. The presence or absence of responses to auditory conditioned stimulation was dependent upon the signal meaning of the stimulus, its physical parameters, and the degree of excitability of the animal.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 4, pp. 539–550, July–August, 1985.     

Neuronal responses in the cat substantia innominata during instrumental conditioned reflex

Neuronal spike activity in the substantia innominata was investigated when establishing the instrumental conditioned reflex of lever-pressing with food reinforcement during chronic experiments on cats. Spike activity changed in 44.3% of neurons during this process; 24.3% responded directly to conditioned acoustic stimulation. Reactions associated with conditioned reflex movement were less pronounced. Response of peak intensity and duration was observed during food reinforcement, when the reaction occasionally occurred 500 msec in advance of the reinforcement provided. Involvement of cholinergic substantia innominata neurons in reinforcement is thought to be responsible for the persistence of residual effects in neocortical neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 6, pp. 796–804, November–December, 1989.     

Avoidance reaction and defensive reflex in the rabbit based on hypothalamic stimulation inducing strokes of the paws

In experiments on rabbits electrical stimulation of the medial hypothalamic area elicited a biphasic reaction: running during the current action and species-specific reaction of kicking the floor with the hind limbs in poststimulous period. Conditioning with acoustic stimulus led in the half of rabbits to elaboration of a habit of jumping on a safe platform in experimental model of active avoidance and to a weak conditioned reaction of kicking without running in the model of classical defensive reflex. The number of conditioned limb kicks was 7-10 times less than that of analogous responses to the reinforcing stimulus. With the increase of the number of combinations, the conditioned habit of kicking became deteriorated: the response became weaker and its latency more prolonged. The results of experiments are discussed in the light of Anokhin's theory of functional system of goal-directed behavioural act.     

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.     

Changes in dry weight of pyramidal cell nuclei in the motor cortex during local motor-food conditioning

The formation of a local motor-food conditioned reflex in rats, consisting of pressing on a level with the unpreferred paw, in one session led to a decrease in dry weight of the large pyramidal cell nuclei in the area of representation of that paw in the motor cortex. Frequent repetition of the local motor-food conditioned reflex consisting of pressing on the level with the preferred paw was not accompanied by any such changes. It is suggested that changes in dry weight of the large pyramidal cell nuclei in the area of representation of the unpreferred paw are due to involvement of these neurons in the formation of the new motor coordination (the local conditioned reflex).     

Intralimbic evoked potentials in the formation of an instrumental food conditioned reflex to hippocampal electrostimulation in dogs]

In dogs with the electrodes implanted in the hippocampus, amygdala, septum and hypothalamus an instrumental alimentary conditioned reflex (CR) was elaborated to electrostimulation of the hippocampus. Intralimbic evoked potentials (EPs) were studied during the elaboration and extinction of this reflex and during stimulations of limbic structures conducted with the purpose of CR generalization checking. Late EP components in the lateral hypothalamus and central nucleus of the amygdala changed during CR elaboration and extinction and in the hippocampus during amygdala testing. In both cases the amplitude of trace positivity and of slow negative wave was less, when during stimulation of the structure an instrumental movement was initiated than at its absence.     

Functional assymmetry of the cerebral cortex in cats during formation of a conditioned reflex

During elaboration of a symmetric active defensive reflex, the animal is given the opportunity to make instrumental movements with the right or left paw. On the first day of elaboration, the cats can be subdivided into three approximately equal groups: with the right preferred paw, with the left preferred paw and ambidexters. After stabilization of the reflex, motor asymmetry increases in both preferred groups, while the number of ambidexters diminishes. Unilateral electric shock producing seizures, which is applied to the "dominant" or "subdominant" hemisphere, exerts correspondingly differing influences on the subsequent reproduction of the reflex. Suppression of reproduction is more pronounced when the shock is applied to the "dominant" hemisphere. The result of repetitive application of electroshock to one and the same hemisphere is that the functions of the "dominant" hemisphere are effected by the hemisphere which was not subjected to direct electrical stimulation.     

The effect of ablation of the caudate nucleus on the alimentary zone of the lateral hypothalamus]

A conditioned food-procuring reaction previously elaborated to an acoustic stimulus was reproduced in chronic experiments on six cats by means of direct electrical stimulation of the posterior parts of the lateral hypothalamus. Folloiwng extensive bilateral electrolytic ablation of the caudate nucleus, conditioned food-procuring reaction to the stimulation of the hypothalamus could not be reproduced for 40 to 70 days. The conditioned foor-procuring reflex to the acoustic stimulus disappeared for 14 to 30 days to be subsequently spontaneously restored. After caudatotomy, a diminution of the average amplitude of background oscillations and of evoked potentials to acoustic stimuli was recorced in the examined zones of the lateral hypothalamus. The part played by the caudate nucleus in the processes of alimentary behaviour activation is discussed.     

Plastic changes in the spontaneous activity of pond snail neurons during rhythmic and associated intracellular electrostimulation

In experiments on spontaneously active neurones of the isolated CNS of pond snail, under contingent stimulation with selective auto-reinforcement of interpulse intervals longer than their mean background value, 70 per cent of neurones were capable of adaptive rearrangements of the initial firing frequency, leading to frequency minimization or maximization of influences applied to them. After control rhythmic stimulation, usually no substantial changes of spontaneous activity were observed. The detected phenomena of initial transformation of the spontaneous impulse activity are considered as a cellular analogue of the instrumental conditioned reflex.     

Fromation of forward and reverse connections during elaboration of a classical and instrumental acid-defense conditioned reflex

During elaboration of a classical acid defensive conditioned reflex to passive lifting of a foreleg in dogs, the reinforcing stimulus--acid elicited an extension of the leg via backward connection. At the initial stage of elaboration of an instrumental avoidance reflex, the reinforcing stimulus called forth an extension of the leg, whereas at the stage of a stable reflex it produced a flexion.     

Functional organization of limbic brain structures during conditioned reflex activity in the dog

In experiments on 3 dogs it was shown that instrumental defensive conditioned reflex (CR) to electrical stimulation of the hippocampus is generalized within the limbic system (LS). The rate of generalization depended on the place of testing stimuli in LS. Stimulation of the medial parts of LS at the stage of stabilized initial reflex did not produce the generalized CRs, while they appeared distinctly in response to stimulation of the lateral parts of LS. In the process of elaboration of heterogeneous CRs (defensive and alimentary) to electrical stimulation of the hippocampus, generalized conditioned reactions from some brain structures appeared in both situations to the same degree while from other structures--with considerable differences. These data point to a possible participation in the genesis of the generalized CRs both of signal and reinforcing brain mechanisms.     

Dynamics of single unit activity in the association cortex of waking cats during defensive conditioning

Responses of neurons in association area 5 during defensive conditioning to acoustic stimulation were studied in chronic experiments on cats. As a rule the neurons responded by excitation to presentation of conditioned and unconditioned stimuli. During the conditioned reflex unit responses usually appeared in the first 50 msec after the beginning of acoustic stimulation, i.e., they were connected with the action of the conditioned stimulus and not with manifestations of conditioned-reflex motion. The most significant changes in responses of cortical association units were observed in the initial period of conditioning. During stabilization of the conditioned reflex, responses of some neurons became stabilized, whereas in other neurons the spontaneous activity and intensity of responses increased, and in a third group the response to one of the stimuli disappeared. This last result indicates a switch during conditioning from polysensory unit responses to monosensory specialized responses. Extinctive inhibition was found to consist of a gradual decrease in the level of the spike discharge and its approximation to spontaneous activity, i.e., to be passive in character.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 6, pp. 563–572, November–December, 1978.     

Role of Acetylcholine in the Modulation of Activity of Neocortical Neurons in Awake Animals Performing an Instrumental Conditioned Reflex

We studied modulatory effects of the cholinergic system on the activity of sensorimotor cortex neurons related to realization of an instrumental conditioned placing reflex. Experiments were carried out on awake cats; multibarrel glass microelectrodes were used for extracellular recording of impulse activity of neurons in the sensorimotor cortex and iontophoretic application of synaptically active agents within the recording region. The background and reflex-related activity was recorded in the course of realization of conditioned movements, and then changes of spiking induced by applications of the testing substances were examined. Applications of acetylcholine and carbachol resulted in increases in the intensity of impulse reactions of neocortical neurons evoked by presentation of an acoustic signal and in simultaneous shortening of the response latencies. An agonist of muscarinic receptors, pylocarpine, exerted a similar effect on the evoked activity of sensorimotor cortex neurons. Blockers of muscarinic receptors, atropine and scopolamine, vice versa, sharply suppressed impulse reactions of cortical neurons to afferent stimulation and simultaneously increased latencies of these responses. Applications of an agonist of nicotinic receptors, nicotine, was accompanied by suppression of impulse neuronal responses, an increase in the latency of spike reactions to presentation of a sound signal, and a corresponding increase in the latency of a conditioned motor reaction. In contrast, application of an antagonist of nicotinic receptors, tubocurarine, significantly intensified neuronal spike responses and shortened their latency. The mechanisms underlying the effects of antagonists of membrane muscarinic and nicotinic cholinoreceptors and the role of activation of these receptors in the modulation of activity of pyramidal and non-pyramidal neocortical neurons related to realization of the instrumental motor reflex are discussed.