The late excitatory responses of the motor cortex neurons in the cat to stimulation of the pyramidal tract

Under conditions of partial suppression of GAMKA-dependent cortical inhibition in the motor cortex of anesthetized cats, a weak electrical stimulation of the pyramidal tract evoked the late slow (50-200 ms) excitatory reactions in the motor cortex neurons similar to those previously recorded under the same conditions in response to stimulation of the parietal cortex. This finding favors the proposal that the late excitatory component of the cortico-cortical response reflects the repetitive activation of cortical neurons due to excitation spread via the system of cortical recurrent excitatory collaterals.     

NMDA-dependent and NMDA-independent neuronal processes in the cat motor cortex,disinhibited by bicuculline,during forepaw placement conditioning

Neuronal activity associated with a conditioned forepaw placing reaction was recorded in the cat's motor cortex locally disinhibited by bicuculline spontaneously diffused from the recording pipette. Electrical stimulation of the parieral cortex (area 5) with 3-5 pulses was used as a conditioned stimulus. In both naive and trained cats, adding of APV (NMDA receptor blocker) led to disappearance of the late (30-120 ms) secondary excitatory responses from the pattern of the neuronal reaction to the parietal stimulation recorded in the motor cortex. At the same time, the APV administration did not change the excitatory reactions (recorded, predominantly, in the deep cortical layers) time-locked to the execution of the conditioned movement. The conditioning resulted in a statistically significant increase in the amplitude and duration of the late secondary responses as well as in a shortening of their latency. In some cases (after a long period of training), the late secondary responses to the conditioned stimulus transformed into paroxysmal epileptiform bursts. A hypothesis is discussed that the increase in synaptic strength of the backward horizontal collaterals of layer-II/III pyramidal neurons is responsible for the learning-related changes in the neuronal reactions in the disinhibited motor cortex.     

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.     

The neuronal reactions of the cat motor cortex to electrical stimulation of the ventral tegmental area as a conditioned signal for the reflex of placing the forelimb on a support]

Electrical stimulation (50-100 pulses, 100-500 Hz) of the ventral tegmental area (VTA) in the vicinity of the n. interpeduncularis in the frontal plane AP2-AP4, L1-L2 caused a cat to grab food placed near its mouth. The conditioned forepaw placing reaction was elaborated using food reinforcement and VTA stimulation as a conditioned stimulus. The conditioned reflex, being once established, was repeatedly performed without extinction in the course of up to 250 trials without food reinforcement. Short (5-10 pulses) conditioned VTA stimulation evoked a prolonged (up to 1000 ms or longer) activation of neurons of the motor cortex and caused a substitution of the inhibitory phase of response to stimulation of the parietal cortex in poststimulus interval in 50-200 ms for the late secondary excitatory response.     

Action of opiate peptides and narcotic analgesics on the cerebral cortex

The chronic experiments on freely moving cats have shown that the opiate peptides, FK33--824 (Tyr--D--Ala--Gly--MePhe--Met(o)--ol) and tetrapeptide (Tyr--D--Ala--Gly--Phe--NH2), as well as the narcotic analgesics, morphine, phentanyl and pentazocine in doses close to analgesic ones, suppress the recovery cycles of primary responses (PR) in the second somatosensory and associative zones of the brain cortex, recorded at paired stimulation of the fibres of thalamo-cortical radiation (TCR). In larger doses these agents slightly increase PR recorded at single stimulation of TCR, provoke the convulsive discharges on EEG and motor excitation of the animals. Naloxon eliminates all the mentioned effects of the tested opiate peptides and narcotic analgesics.     

Bicuculline and orientation tuning of neurons of the visual cortex

Orientation tuning (OT) of 68 visual cortex neurons (field 17) was studied in cats under conditions of a GABA-ergic inhibition blockade by microiontophoretic bicuculline applications; the neuronal responses were evoked by flashing light strips. All characteristics of orientational detection in most neurons got worse after the applications. The OT became wider in 76.3% of cases: its mean value increased from 52.7±2.8° to 85.2±4.6°. In 63.6% of cases OT selectivity decreased by one-third, and in 68.5% of neurons the detection quality decreased by 60%, on average. The threshold dose of bicuculline causing the OT extension was injected by the phoretic current of 31.0±4.5 nA, and the optimum effect was reached at 67.1±6.0 nA. The background activity and the response magnitude increased under the bicuculline influence 3.0 and 4.4 times, respectively, compared with the control. A few minutes after the iontophoresis termination, the frequency of neuronal discharges and OT characteristics returned to their initial values. We conclude that the local blocking of intracortical inhibition, which causes disinhibition of afferent inputs from the neighboring cells with different (compared with the recorded cell) preferred orientations, considerably worsens orientational specificity of visual cortex neurons, or even results in a complete loss of such specificity. These data are consistent with the concept that intracortical inhibition plays a leading role in the formation and sharpening of OT in the visual cortex neurons.Neirofiziologiya/Neurophysiology, Vol. 27, No. 1, pp. 54–62, January–February, 1995.     

Responses of cells in the somatosensory cortex of unanaesthetized cats after sinusoidal displacements of single vibrissae

Summary Neurones in the somatosensory cortex of unanaesthetized restrained cats were recorded during single trapezoid and repetitive sinusoidal displacements of single vibrissae. Responses to trapezoid displacements were similar to those described previously in anaesthetized cats (Hellweg et al., 1977).During repetitive mechanical stimulation cortical cells showed adaptive behaviour so that at higher stimulation frequencies the number of cell discharges per stimulus cycle decreased. The ability to follow the repetition of the stimulus at a one to one ratio was lost in the frequency range between 20 Hz and 60 Hz. A few exceptional cells, while not following at a one to one ratio, still showed some periodicities in their response histograms corresponding to repetition rates of up to 100 Hz. In about 10% of the cells nonmonotonic functions between stimulation frequency and response per cycle were found. These nonmonotonic functions as well as the different adaptive behaviour of cells could not be predicted on the basis of their response to trapezoid stimuli.Measurements of the phase differences between stimulus cycle and response peaks during repetitive stimulation showed that both can vary as a function of stimulation frequency. It is discussed whether these findings could be compatible with the concept of phase coding in the somatosensory cortex.     

Magnetic stimulation of the human motor cortex evokes skin sympathetic nerve activity.

Single-pulse magnetic coil stimulation (Cadwell MES 10) over the cranium induces without pain an electric pulse in the underlying cerebral cortex. Stimulation over the motor cortex can elicit a muscle twitch. In 10 subjects, we tested whether motor cortical stimulation could also elicit skin sympathetic nerve activity (SSNA; n = 8) and muscle sympathetic nerve activity (MSNA; n = 5) in the peroneal nerve. Focal motor cortical stimulation predictably elicited bursts of SSNA but not MSNA; with successive stimuli, the SSNA responses did not readily extinguish (94% of discharges to the motor cortex evoked SSNA responses) and had predictable latencies [739 +/- 33 (SE) to 895 +/- 13 ms]. The SSNA responses were similar after stimulation of dominant and nondominant sides. Focal stimulation posterior to the motor cortex elicited extinguishable SSNA responses. In three of six subjects, anterior cortical stimulation evoked SSNA responses similar to those seen with motor cortex stimulation but without detectable movement; in the other subjects, anterior stimulation evoked less SSNA discharge than that seen with motor cortex stimulation. Contrasting with motor cortical stimulation, evoked SSNA responses were more readily extinguished with 1) peripheral stimulation that directly elicited forearm muscle activation accompanied by electromyograms similar to those with motor cortical stimulation; 2) auditory stimulation by the click of the energized coil when off the head; and 3) in preliminary experiments, finger afferent stimulation sufficient to cause tingling. Our findings are consistent with the hypothesis that motor cortex stimulation can cause activation of both alpha-motoneurons and SSNA.     

Neuronal responses in area 7 of the cat cortex during defensive conditioning

Responses of neurons in area 7 of the parietal association cortex during and after formation of a defensive conditioned reflex to sound were recorded in waking cats. Changes in spike responses of the neurons as a result of the onset of conditioned reflex limb movements were observed in 68% of neurons. Spike responses of neurons formed as a result of learning appeared only if conditioned-reflex limb movements appeared, and they were not observed if, for some reason or other, movements were absent after presentation of the positive conditioned stimulus or on extinction of the reflex. Responses of 46% neurons to conditioned stimulation preceded the conditioned-reflex motor responses by 50–450 msec. The remaining responding neurons were recruited into the response after the beginning of movement. Characteristic spike responses of neurons to the conditioned stimulus appeared 500–900 msec before the beginning of movement and, in the case of appearance of special, "prolonged" motor responses of limb withdrawal, evoked by subsequent reinforcing stimulation.     

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.     

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.     

Effects of repetitive stimulation of the locus coeruleus on spinal inhibitory responses to suprasegmental stimulation in cats

Effects of repetitive stimulation of the locus coeruleus on spinal responses to activation of cortico-, reticulo-, and vestibulospinal tracts were studied in decerebellate cats anesthetized with chloralose. Descending influences of these structures were assessed from changes in amplitude of extensor and flexor monosynaptic discharges or from the magnitude of postsynaptic potentials recorded from the corresponding motoneurons. Stimulation of the motor cortex or modullary reticular formation as a rule evoked two-component inhibitory responses in extensor motoneurons and excitatory-inhibitory responses in flexor motoneurons. Stimulation of locus coeruleus effectively depressed the amplitude of the late component and, to a lesser degree, that of the early component of inhibition arising after stimulation of the cerebral cortex or reticular formation. During stimulation of the locus coeruleus no marked changes were found in inhibitory responses evoked by vestibulospinal influences in flexor motoneurons, and also in excitatory responses arising after stimulation of the above-mentioned descending pathways in both groups of motoneurons.     

Response of cat auditory cortex neurons to tonal stimulation during and after nembutal anesthesia

During acute experiments on 20 cats a comparative study was made of neuronal reaction to a tone, as recorded during the first few hours after administration of Nembutal and after an interval of 10–30 h. No spontaneous activity was seen in 89% of auditory cortex neurons of the anesthetized cats; these produced a sterotyped on- response to the optimal frequency tone. Late neuronal spike discharges at distinct intervals of 100–150 msec appeared in response to the setting up of acoustic stimulation after a brief latent reaction lasting 9–15 msec. It was shown that this stimulation did not produce an off-response in the cortical neurons. When the animals emerged from Nembutal anesthesia, the neurons reacted very differently to the optimal frequency tone. About 76% of the cells produced an on, on-off or off response, while about 21% responded with either tonic spike discharges or total inhibition of these throughout the acoustic stimulation. In unanesthetized cats the vast majority of AI cortical neurons were capable of reacting as long as the stimulus lasted. It is shown how this ability is lost under deep Nembutal anesthetic.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 6, pp. 728–737, November–December, 1985.     

Synaptic effects induced in efferent neurons of the parietal associative cortex of the cat by stimulation of the cerebellar nuclei

Properties of EPSP, evoked in efferent neurons of the parietal associative cortex by stimulation of the cerebellar nuclei, were studied in acute experiments on anesthetized and immobilized cats; intracellular recording was used. The neurons were identified by their antidromic activation after stimulation of the motor cortex, pontinen.n. proprii, or red nucleus. The effects of stimulation of the cerebellar nuclei were of oligo- and polysynaptic nature. The latencies of cerebellofugal EPSP correlated with the latencies of antidromic activation, and correlations were significant both in the cases when the effects of stimulation of separate efferent projections (cortico-cortical, cortico-pontine, or cortico-rubral) and the effects of stimulation of separate cerebellar nuclei were analyzed. The functional role of the efferent systems of the parietal associative cortex and significance of functional parameters of the neurons constituting these systems are discussed.Neirofiziologiya/Neurophysiology, Vol. 27, No. 3, pp. 190–198, May–June, 1995.     

Evoked impulse activity in neuronally isolated cortex

The reactions of neurons of the isolated cortex of one hemisphere to direct cortical stimulation were investigated in cats under Nembutal anesthesia. Isolation of the cortex was carried out by Khananashvili's method [10]. It is shown that phasic reactions develop in the isolated cortex in response to such stimulation: initial discharge, initial pause, first after-discharge, first after-pause, late after-discharges and pauses, as well as reactions of presumably inhibitory neurons. A majority of the cells (85%) which manifest background activity respond to direct electrical stimulation, and the frequency of the late after-reactions is twice as great as in the intact cortex. It is concluded that cortical elements of the isolated cortex retain their principal neurophysiological properties.Institute of Experimental Medicine, Academy of Medical Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 3, No. 3, pp. 236–244, May–June, 1971.     

Distribution of specific signals of different modality in the associative parietal area of the cat cortex

In cats under nembutal anesthesia eliciting specific early components of association responses, the drug parietal distribution upon forepaw and thalamic stimulation was studied: relay somatic-ventrobasal complex (VB) and association nuclei, transmitting specific visual impulses in pulvinar (Pul) and lateral-posterior (LP) areas. Signals of maximum intensity were observed in response to peripheral and central stimulation near somatic area and in response to Pul and LP stimulation in the medial part of parietal cortex. Besides, a general principle revealing more intensive signals of different modality in the areas near lateral sulcus than in other parietal areas was established. The difference in processing of specific polysensory signals in various parietal areas and consequently, different involvement of the latter into the systemic action of the brain was proved, this being related to the character of topical organization of these signals.     

Response activity of different areas of the associative region of the cerebral cortex in the cat

In cats we have simultaneously (monopolarly) recorded the responses in different parts of the associative cortex (ac) (motor cortex, proreal, orbital, anterior marginal, and mid-part of the suprasylvian gyri) appearing for different peripheral stimulations (stimulation of the skin of the forelimbs, a light flash and a sound click). In all the regions of the ac associative responses (ar) almost identical in configuration appeared to all the peripheral stimulations. The ar of the orbitofrontal and the motor cortex differed from the ar of the suprasylvian gyrus in the shorter latent period and greater stability. In each part of the ac we found the same focus of maximum activity for all the peripheral stimulations. For paired stimulations of the same and different modalities the greatest stability in relation to the blocking influence due to the conditioning stimulation characterized the ar which appeared in response to skin stimulation in the orbitofrontal cortex and the ar which appeared in the suprasylvian gyrus on exposure to a light flash. It is assumed that in the orbitofrontal cortex an efferent discharge is formed in response to pulses of different sensory modality whereas in the suprasylvian gyrus there is only sensory integration. Some aspects of afferent convergence are discussed.Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 2, No. 2, pp. 126–139, March–April, 1970.     

The influence of the auditory cortex on acoustically evoked cerebellar responses in the CF-FM bat,Rhinolophus pearsonic chinesis

1. Acoustically evoked responses of 284 neurons isolated from the cerebellar vermis, hemispheres and paraflocculus of Rhinolophus pearsonic chinesis were studied under free field acoustic stimulation conditions. 2. The BFs of these cerebellar auditory neurons ranged from 24 to 76 kHz but they mostly fall either between 48 and 64 kHz or between 65 and 76 kHz. However, the BF distribution varies among vermal, hemispheric and parafloccular neurons. 3. Threshold curves of cerebellar neurons are generally broad but those tuned to the frequency of the predominant CF component are extremely narrow. 4. Response latencies of cerebellar neurons ranged from 2 to 48 ms suggesting multiple auditory cerebellar pathways. The latency distribution also varies among vermal, hemispheric and parafloccular neurons. 5. Although both the vermis and hemispheres contain a disproportionate number of 65-74 kHz neurons, the response latencies of those neurons isolated from the vermis are scattered over a wide range of 2.2-28 ms while those neurons isolated from the hemispheres are generally stabilized between 5 and 12 ms. 6. Electrical stimulation of the auditory cortex evokes discharges from a recorded cerebellar auditory neuron. Cortical stimulation also facilitates the response of an acoustically evoked cerebellar neuron by increasing its number of impulses. The degree of facilitation is dependent upon the amplitude of the acoustic stimulus. 7. For a given electrical and acoustic stimulation condition, the facilitative latency and the degree of facilitation varied with the interstimulus interval. Among 23 neurons studied, most of them (19 neurons, 82.6%) had a maximal facilitative latency between 2 and 10 ms. 8. By examining the difference in the facilitative effect in each isolated cerebellar auditory neuron before and after a topical application of local anesthetic, procaine, onto the point of electrical stimulation in the auditory cortex, we found that the facilitative pathways to vermal and hemispheric neurons may be different from the pathway to parafloccular neurons. 9. Possible auditory pathways to different parts of the cerebellum are discussed in relation to the wide range of recorded response latencies. 10. The facilitative influence of the auditory cortex on the cerebellar auditory neurons is assumed to enhance the cerebellar role in acoustic motor orientation.     

Task-dependent interaction between parietal and contralateral primary motor cortex during explicit versus implicit motor imagery

Both mental rotation (MR) and motor imagery (MI) involve an internalization of movement within motor and parietal cortex. Transcranial magnetic stimulation (TMS) techniques allow for a task-dependent investigation of the interhemispheric interaction between these areas. We used image-guided dual-coil TMS to investigate interactions between right inferior parietal lobe (rIPL) and left primary motor cortex (M1) in 11 healthy participants. They performed MI (right index-thumb pinching in time with a 1 Hz metronome) or hand MR tasks, while motor evoked potentials (MEPs) were recorded from right first dorsal interosseous. At rest, rIPL conditioning 6 ms prior to M1 stimulation facilitated MEPs in all participants, whereas this facilitation was abolished during MR. While rIPL conditioning 12 ms prior to M1 stimulation had no effect on MEPs at rest, it suppressed corticomotor excitability during MI. These results support the idea that rIPL forms part of a distinct inhibitory network that may prevent unwanted movement during imagery tasks.     

Disturbance and restoration of functions after division of spinal afferent pathways in the cat

The possibility and degree of recovery of motor and sensory functions in cats were studied after one-stage or two-stage bilateral division of the posterior columns and spinocervical tracts at the cervical level. Blocking the afferent inflow along these systems led to severe and prolonged disturbances of sensation and motor activity and was accompanied by a sharp decrease in nociceptive sensation. Weak (6–8 V) electrical stimulation of the skin of the limbs, which evoked a primary response of maximal amplitude in intact waking animals, evoked no electrical response in the somatosensory cortex of the chordotomized animals. However, on increasing the intensity of stimulation by 2, 3, or more times, low-amplitude negative waves with a spike latency of about 15 msec, together with slow late waves, were recorded in foci of maximal activity of the cortex. Recovery of motor activity and, to some extent, of proprioception was observed 2–4 months after injury; responses to tactile stimulation were not restored. In the course of compensatory reconstruction evoked activity in the somatosensory cortex did not recover. It is concluded that the recovery of motor activity in cats after injury to the afferent systems of the spinal cord can take place despite a considerable defect of somatic sensation.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 5, No. 3, pp. 281–288, May–June, 1973.