Responses of somatic cortical neurons during the conditioned placing reflex in cats

Unit activity was studied in areas 3 and 4 during the conditioned placing reflex in cats. Responses of somatic cortical neurons in this case were shown to develop comparatively late — 80–100 or, more often, 200–450 msec after the conditioned stimulus. In the motor cortex responses preceded movement by 50–550 msec, whereas in the somatosensory cortex they usually began simultaneously with or after the beginning of the movement. Judging from responses of somatic cortical neurons, the placing reflex is realized by the same neuronal mechanism as the corresponding voluntary movement. The differential stimulus and positive conditioned stimulus, after extinction of the conditioned placing reflex, evoked short-latency spike responses lasting 250–350 msec in the same neurons as took part in the reflex itself. In these types of internal inhibition, responses of the neurons were thus initially excitatory in character. Participation of the neurons in the conditioned placing reflex and its extinction, disinhibition, and differentiation, is the result of a change in the time course of excitatory processes and is evidently connected with differential changes in the efficiency of the various synaptic inputs of the neuron.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 14, No. 4, pp. 392–401, July–August, 1982.     

Effect of response inhibition in cat motor cortex neurons during the expectation of conditioning stimulus

The spike responses of the motor cortex neurons (area 4) associated with forelimb movement were studied in awake cats earlier trained to perform placing motor reactions. Responses produced by the same neurons were compared in two situations: 1) when a sound-click conditioning stimulus (CS) was applied in isolation; 2) when a CS followed a preliminary warning stimulus (WS), a light flash, with a 100–1000 msec delay. During the reflex initiation by combined action of the WS and CS, response components that occurred prior to the placing movement (PM) performance under isolated CS action weakened and arrived 50–150 msec later; yet, response components that appeared in the same situation simultaneously with PM onset or later remained unchanged. PM latent periods were not changed when WS was applied. The temporal interval between WS and CS was characterized by depression of neuronal activity; depression duration was determined by the interstimulus delay. It is conceivable that the described transformations in spike responses of cortical neurons occurred due to changes in the sensory direction of the animal's attention; this direction, in all cases, is a crucial factor in the formation of neuronal activity in the cortex.Translated from Neirofiziologiya, Vol. 25, No. 1, pp. 21–27, January–February, 1993.t     

Neuronal responses of the reticular and anterior ventral thalamic nuclei to stimulation of the thalamic ventrolateral nucleus and motor cortex

Responses of 137 neurons of the rostral pole of the reticular and anterior ventral thalamic nuclei to electrical stimulation of the ventrolateral nucleus and motor cortex were studied in 17 cats immobilized with D-tubocurarine. The number of neurons responding antidromically to stimulation of the ventrolateral nucleus was 10.5% of all cells tested (latent period of response 0.7–3.0 msec), whereas to stimulation of the motor cortex it was 11.0% (latent period of response 0.4–4.0 msec). Neurons with a dividing axon, one branch of which terminated in the thalamic ventrolateral nuclei, the other in the motor cortex, were found. Orthodromic excitation was observed in 78.9% of neurons tested during stimulation of the ventrolateral nucleus and in 52.5% of neurons during stimulation of the motor cortex. Altogether 55.6% of cells responded to stimulation of the ventrolateral nucleus with a discharge of 3 to 20 action potentials with a frequency of 130–350 Hz. Similar discharges in response to stimulation of the motor cortex were observed in 30.5% of neurons tested. An inhibitory response was recorded in only 6.8% of cells. Convergence of influences from the thalamic ventrolateral nucleus and motor cortex was observed in 55.7% of neurons. The corticofugal influence of the motor cortex on responses arising in these cells to testing stimulation of the ventrolateral nucleus could be either inhibitory or facilitatory.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 5, pp. 460–468, September–October, 1978.     

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.     

Transformation of the afferent tactile signal into a motor command in the cat motor cortex

Activity of 112 neurons of the precruciate motor cortex in cats was studied during a forelimb placing reaction to tactile stimulation of its distal parts. The latent period of response of the limb to tactile stimulation was: for flexors of the elbow (biceps brachii) 30–40 msec, for the earliest reponses of cortical motor neurons about 20 msec. The biceps response was observed 5–10 msec after the end of stimulation of the cortex with a series of pulses lasting 25 msec. Two types of excitatory responses of the neurons were identified: responses of sensory type observed to each tactile stimulation of the limb and independent of the presence or absence of motion, and responses of motor type, which developed parallel with the motor response of the limb and were not observed in the absence of motion. The minimal latent period of the responses of motor type was equal to the latent period of the sensory responses to tactile stimulation (20±10 msec). Stimulation of the cortex through the recording microelectrode at the site of derivation of unit activity, which increased during active flexion of the forelimb at the elbow (11 stimuli at intervals of 2.5 msec, current not exceeding 25 µA), in 70% of cases evoked an electrical response in the flexor muscle of the elbow.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 9, No. 2, pp. 115–123, March–April, 1977.     

Comparison of neuronal response pattern in the rat somatosensory cortex during active and passive vibrissae movements

A comparative study of neuronal response in separate cortical columns of the somatosensory cortex (the barrel field area) was made in unanesthetized partially curarized white rats under various circumstances: during passive deflection of immobile vibrissa, unhindered volitional sweeping movement of the vibrissae, and during movement induced by stimulating the motor cortex and facial muscles. Differences in the response of the same neurons emerged under these different experimental situations. Different groups of neurons — responding before, during, and after volitional vibrissa movements were observed. Such response is thought to be triggered by different afferent trains reaching cortical column neurons from sources including the motor cortex, the vibrissa follicle receptors, and facial muscles.Institute of Neurocybernetics, State University, Rostov-on-Don. State University, Simferopol. Translated from Neirofiziologiya, Vol. 22, No. 2, pp. 235–242, March–April, 1990.     

Dynamics of cortical unit responses during defensive conditioning to sound

Several phases were distinguished in single-unit responses in areas 3 and 4 during defensive conditioning to acoustic stimulation: an initial response, short inhibition of the spike discharge, early and late after-discharges, and changes arising after the end of acoustic stimulation. The initial spike response appeared or intensified (if present already) in the first period of defensive conditioning parallel with an increase in spontaneous unit activity. After-discharges appeared later. The conditioned-reflex movement usually began 100–400 msec after stimulation began. This latent period of the first movement was the same whether for a real conditioned reflex or an after-discharge. Comparison of the latent periods of conditioned movements with the phases of the unit responses showed that the conditioned responses of the cortical neuron were primarily modified after-discharges of neurons evoked by a conditioned stimulus. Differential unit responses to acoustic stimulation, also based on after-discharges, were formed just as actively as positive. The basic role of reinforcement during conditioning is not to increase the excitability of the neurons, which is important in connection with their acquisition of polysensory properties, but to modify the after-discharges of the neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 4, pp. 339–347, July–August, 1978.     

Responses of cat pulvinar neurons to moving visual stimuli

Responses of 114 pulvinar neurons to moving visual stimuli were studied. Most (79) neurons possessed spontaneous activity (10–25 spikes/sec). Of 59 neurons tested, 31 responded to stimulation of both retinas and 28 to stimulation only of the contralateral retina. Of 114 neurons, 41 responded only to movement of black objects, while the rest responded to movement of both black and light objects. According to the character of their responses to movement the neurons were divided into two main groups. The first group consisted of neurons sensitive to the direction of motion and responding with a spike discharge to movement in one direction and by inhibition to movement in the opposite direction. The second group included neurons insensitive to the direction of motion and responding by an equal number of discharges to movements in two opposite directions. Besides these two main groups, other neurons responding to movement in two opposite directions by discharges with different temporal distribution and also neurons which changed the character of their response from nondirectional to directional depending on the size of the moving stimulus, were found.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 10, No. 4, pp. 348–354, July–August, 1978.     

Sensory input to neurons of the motor projection of biceps in the cat cortex

Correlation of cortical unit activity in the motor area for the biceps muscle was studied in chronic experiments on cats. In a group of neurons whose activity correlated with movement 68.1% of units had no sensory input from the working limb whereas in a group of neurons not correlating with movement there were 97.6% such units. In 24.2% of group I neurons cutaneous receptive fields of activation type were discovered on the distal part of the dorsal surface of the working limb. Five neurons responding to sensory input from the joints of the working limb were studied in this group.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 9, No. 6, pp. 563–569, November–December, 1977.     

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.     

Unit responses of the secondary somatosensory cortex during defensive conditioning in cats

Unit responses in the secondary somatosensory cortex during the formation and extinction of a defensive conditioned reflex to acoustic stimulation were investigated in chronic experiments on cats. In 21 of 28 neurons tested during defensive conditioning the firing pattern changed in accordance with the character of responses to electric shock reinforcement. Two types of conditioned-reflex unit responses were distinguished: excitatory and inhibitory. Most neurons responding to the conditioned stimulus by activation did so during the first 50 msec, which was 80–100 msec before the conditioned motor response. Considerable variability of the unit responses was observed during conditioning. By the time of stabilization of the conditioned-reflex connections the unit response to the conditioned stimulus was stable in form. The pattern of extinction of the conditioned unit activity was expressed as a decrease in the discharge frequency in responses of excitatory type and disinhibition of activity in the case of inhibitory responses.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev, Translated from Neirofiziologiya, Vol. 9, No. 3, pp. 232–238, May–June, 1977.     

Responses of secondary sensomotor cortical neurons to electrodermal and acoustic stimulation in waking cats

Unit responses in the second somatosensory cortical projection area (SII) to clicks and electric shocks applied to the contralateral limb were investigated in chronic experiments on cats. In response to specific stimulation for the cortical region studied the discharge frequency of 75% of neurons increased, spontaneous activity of 18% was reduced in frequency or the discharges ceased altogether, and 25% of cells did not respond. In response to "nonspecific" stimulation (clicks) 30% of neurons were activated; the discharge of 25% of cells was inhibited and 45% did not respond. The results of investigation of intersensory convergence of stimuli from different sensory systems showed that a high proportion (55%) of SII neurons give bimodal responses. Another 18% of neurons give a specific response to both adequate and inadequate stimulation. It is suggested that the presence of polysensory convergence of SII neurons and of short pathways for the conduction of sensory information, and also the ability of neurons to acquire polysensory properties during stimulus presentation are evidence of the important role of this cortical region in conditioning.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 9, No. 5, pp. 453–459, September–October, 1977.     

Two patterns of bulbar neuronal response to microstimulation of locomotor and inhibitory brain stem sites

Synaptic responses (postsynaptic potentials and action potentials) were evoked in mesencephalic decerebellated cats by stimulating pontine bulbar locomotor and inhibitory sites (LS and IS, respectively) with a current of not more than 20 µA in "medial" and "lateral" neurons of the medulla. Some neurons even produced a response to presentation of single (actually low — 2–5 Hz — frequency) stimuli. The remaining cells responded to stimulation at a steady rate of 30–60 Hz only. Both groups of medial neurons were more receptive to input from LS. Lateral neurons responding to even single stimuli reacted more commonly to input from LS and those responding to steady stimulation only to input from IS. Many neurons with background activity (whether lateral or medial) produced no stimulus-bound response, but rhythmic stimulation either intensified or inhibited such activity. This response occurs most commonly with LS stimulation. Partial redistribution of target neurons in step with increasing rate of presynaptic input may play a major part in control of motor activity.Institute for Research into Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 22, No. 2, pp. 257–266, March–April, 1990.     

Responses of limbic cortical neurons during instrumental conditioned reflexes in cats

Unit activity in cortical areas 24 and 32 was studied during conditioned placing reflex formation in cats. Neuronal responses in the limbic cortex of trained animals correlated with acoustic stimulation, the motor response, and also with the presentation of food reinforcement. In untrained animals 16% of neurons responded to acoustic stimulation. After training the number of neurons responding to sound in area 32 increased to 51.3%. Of the total number of neurons, 34.6% responded by initial excitation and 26.7% by inhibition of spike activity. The latent period of these responses was about 50 msec and their duration up to 200 msec. Similar but weaker responses were observed in area 24. Short-latency activation responses to conditioned and differential stimulation were similar in character. It is suggested that after training processes taking place in the limbic cortex may contribute to better perception of both conditioned and differential acoustic stimuli, irrespective of their functional significance.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 16, No. 2, pp. 201–208, March–April, 1984.     

Cross-correlation analysis of unit activity in spinal locomotor centers

Correlation analysis of unit activity in spinal locomotor centers was carried out in immobilized thalamic cats. Within a short time interval (the time shift of one spike train relative to the other during plotting of the cross-correlation histogram did not exceed 54 msec) correlation between the spike flows of these cells was absent, irrespective of the distance between them, both at rest and during efferent discharge generation. Spike flows of neurons could correlate only in the case of a long time interval (maximal time shift of one spike train relative to the other not less than 4–8 sec during plotting of the cross-correlation histogram). Weak correlation with a long time interval (4–8 sec) was found between changes in the momentary frequency of a neuron and the intensity of the discharge in the motor nerve, but no correlation was found between changes in momentary frequency of the neuron and intensity of discharge. The possible causes of the absence of correlation with a short time interval and its presence with a long time interval are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 3, pp. 283–289, May–June, 1980.     

Neurons of the cat sensorimotor cortex: Responses related to differentiation of monomodal and heteromodal conditioning signals

Spike responses of the sensorimotor cortex neurons were studied in chronic experiments on cats trained to perform an operant reflex, a placing movement triggered by application of the distant stimuli. The responses recorded under conditions of differentiation of sound tones of various frequencies and of a heteromodal complex (light + tone) from its components were compared. The responses recorded from 125 neurons in 3 animals were analyzed. No neurons selectively responding either to monomodal or to heteromodal signals were found. Forty-five cells responded to positive signals by excitation or inhibition, irrespective of the signal modality. The shortest latencies of these responses were 30 and 40 msec, respectively. When inhibitory stimuli were applied, these neurons either generated much weaker responses, or did not respond at all. A correlation was found between the level of response depression and the level of differentiation of the signals by an animal. These findings allow us to hypothesize that the sensorimotor cortex does participate in differentiation of sensory signals, providing preparation for switching on the motor response after a positive stimulus or suppression of such a response after a negative one.Neirofiziologiya/Neurophysiology, Vol. 26, No. 4, pp. 251–261, July–August, 1994.     

Influence of food motivation on neuronal response of the cat somatic cortex during instrumental reflex

Spike activity in neurons of areas 3 and 4 was investigated in experiments on cats during the conditioned reflex response of placing the paw on a support both before and after feeding ad libitum. Ingestion of a feed consisting of a rapidly absorbed glucose dairy mix did not prevent the reflex taking place if the animals' favorite food was used as reinforcement. Background activity increased in two-thirds of the neurons after the feed; the tonic constituent of neuronal response declined substantially and repeated contraction of the biceps occurring at the same rate as locomotor movements disappeared. Difference in latency of response produced by the conditioned stimulus in the same neurons before and after feeding measured 50–300 msec during the experiment. Measurements of latency of placing motion remained largely unchanged. Changes in the latency of neuronal spike response were thus found to be interrelated with the intensity of the animal's motivational excitation. It is suggested that fluctuations in degree of food motivation lead to changes in cortico-subcortical relationships responsible for initiation and performance of conditioned movements in these animals.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 19, No. 5, pp. 646–653, September–October, 1987.     

Somato-Motor Haptic Processing in Posterior Inner Perisylvian Region (SII/pIC) of the Macaque Monkey

The posterior inner perisylvian region including the secondary somatosensory cortex (area SII) and the adjacent region of posterior insular cortex (pIC) has been implicated in haptic processing by integrating somato-motor information during hand-manipulation, both in humans and in non-human primates. However, motor-related properties during hand-manipulation are still largely unknown. To investigate a motor-related activity in the hand region of SII/pIC, two macaque monkeys were trained to perform a hand-manipulation task, requiring 3 different grip types (precision grip, finger exploration, side grip) both in light and in dark conditions. Our results showed that 70% (n = 33/48) of task related neurons within SII/pIC were only activated during monkeys’ active hand-manipulation. Of those 33 neurons, 15 (45%) began to discharge before hand-target contact, while the remaining neurons were tonically active after contact. Thirty-percent (n = 15/48) of studied neurons responded to both passive somatosensory stimulation and to the motor task. A consistent percentage of task-related neurons in SII/pIC was selectively activated during finger exploration (FE) and precision grasping (PG) execution, suggesting they play a pivotal role in control skilled finger movements. Furthermore, hand-manipulation-related neurons also responded when visual feedback was absent in the dark. Altogether, our results suggest that somato-motor neurons in SII/pIC likely contribute to haptic processing from the initial to the final phase of grasping and object manipulation. Such motor-related activity could also provide the somato-motor binding principle enabling the translation of diachronic somatosensory inputs into a coherent image of the explored object.     

Changes in the activity of ventrolateral thalamic neurons and instrumental response after systemic administration of N-methyl-4-phenyl-1,2,3,6-tetrahydropyridine (MPTP)

Neuronal activity in the ventrolateral thalamus during execution of instrumental reaction before and after parenteral administration of N-methyl-4-phenyl-1,2,3,6-tetrahydropyridine (MPTP) was investigated in samples of 81 and 70 cells, respectively. After a 5-day course of one 5 mg/kg MPTP injection daily, firing rate of neurons in which activity correlated with forelimb movement rose significantly; this activation increased in length during the initial, flexor, and extensor stages of motor response. Bradykinesia set in together with intensified neuronal activation in the animals. Microinjection of exogenous dopamine into the caudate nucleus brought about correction of motor disturbance and a reduced neuronal firing rate in the ventrolateral (thalamic) nucleus. It was deduced that the nigrostriatal system exercises inhibitory control over the activity of thalamic neurons associated with forelimb movement in thalamic neurons in intact animals.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 22, No. 3, pp. 291–300, May–June, 1990.     

Temporal summation in cat visual cortical neurons

Characteristics of temporal summation in neurons of area 17 of the visual cortex in acute experiments on unanesthetized, immobilized cats. During light adaptation, extracellular spike responses of these neurons to optimal local photic stimuli of varied duration — from 5 to 1000 msec — were studied. The critical duration of temporal summation of excitation, determined by the supraliminal method using the criterion of maximal discharge frequency in the first volley of the spike response, varied in different cells from 5 to 100 msec; neurons with summation lasting 15–100 msec (mean 31.45±5.67 msec) were found most frequently. Neurons with central receptive fields differed significantly from cellswith peripheral fields in the shorter critical duration of temporal summation, the lower frequency of spontaneous discharges, and the shorter duration of the first volley of the response. Summation time in neurons with simple receptive fields was significantly shorter than in neurons with complex receptive fields. The results of these experiments are compared with data in the literature obtained by the study of retinal and lateral geniculate neurons in cats and are discussed from the stand-point of division of ascending afferent projections in the visual system into X-and Y-groups (Ia and Ib).Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 13, No. 4, pp. 345–352, July–August, 1981.