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.     

Neuronal responses of a chronically isolated slab of auditory cortex to intracortical stimulation during paroxysmal electrical activity in cats

Responses of 155 neurons 3 weeks after neuronal isolation of a slab of auditory cortex (area AI) to single intracortical stimulating pulses at the level of layer IV were studied in unanesthetized, curarized cats during paroxysmal electrical activity evoked by series of high-frequency (10–20 Hz) electrical stimulation by a current 2–5 times above threshold for the direct cortical response. In response to such stimulation a discharge of paroxysmal electrical activity, lasting from a few seconds to tens of seconds, appeared in the slab. As a rule it consisted of two phases — tonic and clonic. This indicates that cortical neurons can form both phases of paroxysmal cortical activity. Depending on behavior of the neurons during paroxysmal electrical activity and preservation of their ability to respond to intracortical stimulation at this time, all cells tested in the isolated slab were divided into four groups. Their distribution layer by layer and by duration of latent periods was studied. Two-thirds of the neurons tested were shown to generate spike activity during paroxysmal discharges whereas the rest exhibited no such activity. A special role of neurons in layer II in generation of paroxysmal activity in the isolated slab was noted. The view is expressed that at each moment functional neuronal circuits, independent of each other, exist in the slab and also, evidently in the intact cortex, which can interact with one another when conditions change.I. I. Mechnikov Odessa State University. Translated from Neirofiziologiya, Vol. 16, No. 1, pp. 3–11, January–February, 1984.     

Monosynaptic inhibitory postsynaptic potentials of cortical neurons

Monopolar intracortical stimulation of the auditory cortex was carried out in cats immobilized with D-tubocurarine. A macroelectrode (tip diameter 100 µ) or a microelectrode (tip diameter 10–15 µ) was used for stimulation. In both cases, besides excitatory responses, primary IPSPs with latent periods of 0.4–1.2 and 1.4–6.0 msec were recorded in cortical neurons close to the point of stimulation. The first group of IPSPs are considered to be generated in response to direct stimulation of bodies or axons of inhibitory cortical neurons, i.e., monosynaptically. The amplitude of these IPSPs varied in different neurons from 3 to 15 mV, and their duration from 4 to 150 msec. Additional later inhibitory responses were superposed on many of them. Of the IPSPs generated in auditory cortical neurons in response to stimulation of geniculocortical fibers 1.5% had a latency of 0.8–1.3 msec. They also are assumed to be monosynaptic. It is concluded that the duration of synaptic delay of IPSPs in cortical neurons and spinal motoneurons is the same, namely 0.3–0.4 msec. Axons of auditory cortical inhibitory neurons may be 1.5 mm long. The velocity of impulse conduction along these axons is 1.6–2.8 m/sec. The genesis of some special features of IPSPs of cortical neurons is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 7, No. 5, pp. 458–467, September–October, 1975.     

Functional characteristics of visual cortical neurons of the cat

The characteristics of neurons in Area 17 of the visual cortex in cats were investigated by extracellular recording of their activity. Unit responses to flashes modulated by intensity and duration (100 µsec-1 sec) were recorded. Of 80 neurons tested, 67.6% were spontaneously active and 32.4% were silent. The threshold responses of the neurons to flashes varied by 7 logarithmic units. The distribution curve of the cells by response thresholds had one maximum corresponding to an energy of the order of 1–10 lm·sec. The time during which the cells could summate excitation did not exceed a mean value of 34 msec. Depending on the latent periods of the visual cortical neurons they can be divided into three groups. The first group includes neurons responding 20–40 msec after stimulation, the second and third neurons responding after 100–120 and 160–180 msec, respectively. Photic stimulation considerably altered the ratio between the numbers of cells generating spikes with high and low frequency. No correlation was found between the sensitivity of the visual cortical cells to light, the latent period of their response, and the critical time of summation. This shows that the cortex contains many duplicate units which are grouped together on the basis of only one of the functional characteristics of their spike response.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 2, No. 2, pp. 173–179, March–April, 1970.     

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.     

Postsynaptic neuronal response of a cat sensorimotor cortex during evoked and self-sustained rhythmic "spike and wave" activity

Intracellular correlates of complex sets of rhythmic cortical "spike and wave" potentials evoked in sensorimotor cortex and of self-sustained rhythmic "spike and wave" activity were examined during acute experiments on cats immobilized by myorelaxants. Rhythmic spike-wave activity was produced by stimulating the thalamic relay (ventroposterolateral) nucleus (VPLN) at the rate of 3 Hz; self-sustained afterdischarges were recorded following 8–14 Hz stimulation of the same nucleus. Components of the spike and wave afterdischarge mainly correspond to the paroxysmal depolarizing shifts of the membrane potential of cortical neurons in length. After cessation of self-sustained spike and wave activity, prolonged hyperpolarization accompanied by inhibition of spike discharges and subsequent reinstatement of background activity was observed in cortical neurons. It is postulated that the negative slow wave of induced spike and wave activity as well as slow negative potentials of direct cortical and primary response reflect IPSP in more deep-lying areas of the cell bodies, while the wave of self-sustained rhythmic activity is due to paroxysmal depolarizing shifts in the membrane potential of cortical neurons.I. S. Beritashvili Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 18, No. 3, pp. 298–306, May–June, 1986.     

Characteristics of electrical activity of olfactory bulb neurons of fish

It was shown by intracellular recording of the activity of olfactory bulb neurons of the carp that their dendrites are excited both by synaptic activation and by direct stimulation with an electric current. The dendrites generate an action potential and probably conduct it for some distance toward the soma. The neurons can be divided into two groups: one responds to ortho- and antidromic stimuli with one, rarely with two peaks, the other responds with a rhythmical discharge. The presence of early and late IPSP is characteristic of neurons of both groups. Rhythmical variations in potential with a frequency of 26–33/sec, so-called oscillations, are recorded; they may be excitatory (in secondary neurons they correspond to EPSP) or inhibitory (they correspond to IPSP). Possible mechanisms of the excitatory oscillations and the rhythmical discharge in olfactory bulb neurons of the carp are discussed.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol.3, No.5, pp. 505–511, September–October, 1971.     

Two types of granular cells in the cerebellar cortex

We recorded the activity of two types of granular cells in the rostral folia of the paramedial lobe (the projection region of the front legs) of the cerebellar cortex in cats immobilized by administration of ditiline; these cells differed in their receptive fields, the characteristics of their reaction to single stimulation of somatic nerves, and the character of their background activity. The granular cells of the first type were excited only when the nerves of the front legs were stimulated (reacting with 1–3 impulses with a latent period of 8–20 msec) and were inhibited between 20–50 and 70–180 msec after stimulation of the nerves of any leg. The cells of the second type responded with volleys of 3–6 impulses with a latent period of 20–40 msec to stimulation of the nerves of all four legs. Comparison of the reactions of the granular cells and other neurons of the cerebellar cortex showed that the cells of the first type cause excitation of the Purkinje and Golgi cells and the neurons of the molecular layer. The granular cells of the second type have an excitatory effect on the Golgi cells. The differences in the reactions of the two types of granular cells result from the fact that they are selectively innervated by the mossy fibers of different afferent pathways. Comparison with the data in the literature enables us to surmise that the fibers of the cuneocerebellar tract terminate at granular cells of the first type, while the reticular fibers terminate at cells of the second type.Institute of Problems of Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 1, No. 2, pp. 167–176, September–October, 1969.     

Activity of sensorimotor cortex neurons during a dopaminergic transmission block in cats performing conditional reflex

Activity of sensorimotor cortex neurons was recorded in chronic experements on cats trained to perform instrumental conditional reflex; records were made before, during, and after isolated iontophoretic applications of haloperidol or glutamate, or their combined application. Haloperidol was shown either to facilitate or to inhibit the background and evoked (related to acoustic stimulation and motor response) spike activity of cortical neurons. Aftereffects of haloperidol were observed too; they were still expressed 10–15 min after the cessation of the iontophoresis. Combined haloperidol and glutamate application was followed by a sharp suppression of the evoked responses potentiated earlier by glutamate. An adenylatecyclase system is supposed to mediate the facilitation evoked by glutamate application. Some modulators, including dopamine, probably activate adenylatecyclase and in this way ensure facilitation of the glutamate-induced responses.Neirofiziologiya/Neurophysiology, Vol. 26, No. 5, pp. 347–355, September–October, 1994.     

Unit responses in the cat auditory cortex to electrical stimulation of nerve fibers innervating receptor cells in different parts of the organ of corti

Responses of 200 primary auditory cortical neurons to electrical stimulation of nerve fibers in different receptor zones of the cochlea were studied in cats anesthetized with pentobarbital. Under the influence of paired stimulation, after the response to the conditioning stimulus a state of prolonged (from 4 to 200 msec) refractiveness to the second stimulus developed in all the neurons tested. This long-lasting inhibition of unit activity was due to inhibition developing in the thalamus and the auditory cortex itself. The intensity and duration of excitation and inhibition in the cortical projection focus were maximal when the center of the receptive field was stimulated and decreased when the stimulus shifted from the center to the periphery. The region of the receptor surface of the cochlea to stimulation of which the auditory cortical neurons respond by an action potential is much narrower than the region whose electrical stimulation depresses the discharge of these neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 14, No. 4, pp. 418–425, July–August, 1982.     

Responses of rabbit visual cortical neurons to intracortical electrical stimulation

Responses of rabbit visual cortical neurons to single and repetitive intracortical electrical stimulation were investigated. The stimulating electrode was located 0.7–1.2 mm away from the recording electrode. Response thresholds to single stimulation were as a rule 150–180 µA, whereas to series of stimuli they were 30–60 µA. The latent period to the first spike averaged 5–15 msec but the probability of the initial discharge was very low, namely 3–6%. With an increase in current intensity the duration of the initial inhibitory pause was increased in half of the neurons responding to it, whereas in the rest it was unchanged. After presentation of series of stimuli spontaneous activity was enhanced for a short time (4–6 sec). In about half of the cells the same kinds of discharge dynamics were observed in response to repetitive stimulation (frequency 0.25 Hz) as in responses to light, but more neurons with sensitization of discharge and fewer "habituating" neurons took part in responses to electrical stimulation. It is postulated that stimulation of a given point of the visual cortex evokes excitation of a local neuron hypercolumn and inhibition of neighboring cell columns.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 15, No. 4, pp. 412–419, July–August, 1983.     

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.     

Responses of parietal associative neurons to stimulation of primary sensory areas

Responses of 251 neurons in the anterior part of the middle suprasylvian gyrus to stimulation of primary sensory (auditory, visual, somatosensory) areas and also to acoustic, visual, and somatosensory stimuli were studied in acute experiments on cats anesthetized with chloralose (40 mg/kg) and pentobarbital (20 mg/kg). Three groups of neurons were distinguished by their responses to stimulation of the primary sensory areas: those responding by an increased firing rate (117) or by inhibition (35) and those not responding (99). Responses of 193 neurons to stimulation of the peripheral afferent systems were analyzed. Neurons of the parietal associative cortex responded more frequently to cortical stimulation than to peripheral. By the duration of the latent period of their response to cortical stimulation the neurons were divided into three groups: those with short (less than 20 msec), medium (20–30 msec), and long latent periods (over 30 msec). The first group was the largest.Kemerovo State Medical Institute. Translated from Neirofiziologiya, Vol. 4, No. 5, pp. 524–530, September–October, 1972.     

Unit responses of the first and second somatosensory cortical areas to peripheral,thalamic, and cortical stimulation

Unit responses of the first (SI) somatosensory area of the cortex to stimulation of the second somatosensory area (SII), the ventral posterior thalamic nucleus, and the contralateral forelimb, and also unit responses in SII evoked by stimulation of SI, the ventral posterior thalamic nucleus, and the contralateral forelimb were investigated in experiments on cats immobilized with D-tubocurarine or Myo-Relaxin (succinylcholine). The results showed a substantially higher percentage of neurons in SII than in SI which responded to an afferent stimulus by excitation brought about through two or more synaptic relays in the cortex. In response to cortical stimulation antidromic and orthodromic responses appeared in SI and SII neurons, confirming the presence of two-way cortico-cortical connections. In both SI and SII intracellular recording revealed in most cases PSPs of similar character and intensity, evoked by stimulation of the cortex and nucleus in the same neuron. Latent periods of orthodromic spike responses to stimulation of nucleus and cortex in 50.5% of SI neurons and 37.1% of SII neurons differed by less than 1.0 msec. In 19.6% of SI and 41.4% of SII neurons the latent period of response to cortical stimulation was 1.6–4.7 msec shorter than the latent period of the response evoked in the same neuron by stimulation of the nucleus. It is concluded from these results that impulses from SI play an important role in the afferent activation of SII neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 4, pp. 351–357, July–August, 1976.     

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.     

Negative surface potential shift and responses of neurons and glial cells during tetanic stimulation of the cortical surface

The negative potential shift in response to tetanic stimulation of the surface of the cortex or thalamic nucleus was recorded from the cortical surface in cats lightly anesthetized with pentobarbital. Parallel intracellular recordings were obtained of activity of neurons and glial cells. Glial cells responded to this stimulation by slow depolarization, which, under certain conditions of stimulation, was followed by slow hyperpolarization; hyperpolarization shifts were observed in neurons. Depolarization and hyperpolarization of glial cells, like hyperpolarization of neurons, did not correlate in time with the development of a negative shift of the surface potential. It is postulated that this shift is a response of complex origin involving the participation not only of glial cells, but also of cortical neurons.I. S. Beritashvili Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 14, No. 3, pp. 248–253, May–June, 1982.     

Responses of medial geniculate body neurons to auditory cortical stimulation

Extracellular and intracellular unit responses of thepars principalis of the medial geniculate body to stimulation of the first (AI), second (AII), and third (AIII) auditory cortical areas were studied in cats immobilized with D-tubocurarine. In response to auditory cortical stimulation both antidromic (45–50%) and orthodromic (50–55%) responses occurred in the geniculate neurons. The latent period of the antidromic responses was 0.3–2.5 msec and of the orthodromic 2.0–18.0 msec. Late responses had a latent period of 30–200 msec. Of all neurons responding antidromically to stimulation of AII, 63% responded antidromically to stimulation of AI also, confirming the hypothesis that many of the same neurons of the medial geniculate body have projections into both auditory areas. Orthodromic responses of geniculate neurons consisted either of 1 or 2 spikes or of volleys of 8–12 spikes with a frequency of 300–600/sec. It is suggested that the volleys of spikes were discharges of inhibitory neurons. Intracellular responses were recorded in the form of antidromic spikes, EPSPs, EPSP-spike, EPSP-spike-IPSP, EPSP-IPSP, and primary IPSP. Over 50% of primary IPSP had a latent period of 2.0–4.0 msec. It is suggested that they arose through the participation of inhibitory interneurons located in the medial geniculate body.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 1, pp. 5–12, January–February, 1976.     

Neuronal response in a strychninized cortical slab isolated from the cat

Intracellular response in neurons and glial cells of an isolated cortical slab to direct electrical stimulation of the slab following surface application of strychnine was investigated during experiments on immobilized unanesthetized cats. Strychnine induced single epileptiform discharges and after-discharges in the slab and in the neurons it contained in the form of large-scale paroxysmal depolarization shifts (PDS) in membrane potential (MP). Spontaneous summated epileptiform discharges and neuronal activity in the units examined were not very synchronized. Electrical stimulation induced generalized paroxysmal activity in the isolated slab. Neuronal PDS were accompanied by refractory periods, onset of which did not depend on MP level. Strychnine increased the number of neurons manifesting background activity in which action potentials were generated by rhythmic depolarizing MP waves of extrasynaptic origin. Epileptiform response in strychninized cortical isolated slabs to presentation of single stimuli is accompanied by major depolarization shifts in the MP of glial cells. Paroxysmal excitation is thought to be triggered in strychninized isolated cortical slabs by extrasynaptic factors and closely linked to altered concentration of extracellular potassium.I. I. Mechnikov University, Odessa. Translated from Neirofiziologiya, Vol. 22, No. 1, pp. 23–29, January–February, 1990.     

Effects of food motivation on spike activity of cat somatic cortex neurons

Spike response to unconditioned electrocutaneous stimulation was investigated in cortical neurons of areas 3 and 4 in untrained hungry cats during heightened excitation motivated by food presentation and when at rest. This reinforcement led to changed background activity level, reduced intensity of the initial stages of spike response, and disappearance of late neuronal response. Neuronal response of the same cortical area to a conditioned stimulus (a clicking sound) during reduced food motivation (the animals being sated during the course of the experiment) was also studied under the effects of instrumental feeding reflex. Coordination between the timing of neuronal response and their corresponding movements was discovered from comparing response pattern accompanying the execution of paw-placing (conditioned reflex and intersignal) movements and those recorded at different levels of food-induced excitation, as well as a similarity between these reactions. It was found that the initial stages of neuronal response to a conditioned signal only occurred during contraction of the brachial biceps muscle, while coordination between their timing and that of EMG changes was dependent on the animal's degree of satiation. Findings indicate the possibility of food-induced excitation substantially influencing spike response pattern in somatic cortex neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 19, No. 6, pp. 725–735, November–December, 1987.     

Neuronal and synaptic mechanisms of cortical inhibition

The latent periods, amplitude, and duration of IPSPs arising in neurons in different parts of the cat cortex in response to afferent stimuli, stimulation of thalamocortical fibers, and intracortical microstimulation are described. The duration of IPSPs evoked in cortical neurons in response to single afferent stimuli varied from 20 to 250 msec (most common frequency 30–60 msec). During intracortical microstimulation of the auditory cortex, IPSPs with a duration of 5–10 msec also appeared. Barbiturates and chloralose increased the duration of the IPSPs to 300–500 msec. The latent period of 73% of IPSPs arising in auditory cortical neurons in response to stimulation of thalamocortical fibers was 1.2 msec longer than the latent period of monosynaptic EPSPs evoked in the same way. It is concluded from these data that inhibition arising in most neurons of cortical projection areas as a result of the arrival of corresponding afferent impulsation is direct afferent inhibition involving the participation of cortical inhibitory interneurons. A mechanism of recurrent inhibition takes part in the development of inhibition in a certain proportion of neurons. IPSPs arise monosynaptically in 2% of cells. A study of responses of cortical neurons to intracortical microstimulation showed that synaptic delay of IPSPs in these cells is 0.3–0.4 msec. The length of axons of inhibitory neurons in layer IV of the auditory cortex reaches 1.5 mm. The velocity of spread of excitation along these axons is 1.6–2.8 msec (mean 2.2 msec).A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 16, No. 3, pp. 394–403, May–June, 1984.