Electrophysiological study of connections between the entorhinal cortex and the neocortex

In acute experiments in rabbits immobilized by d-tubocurarine, stimulation of the entorhinal area with rectangular electric impulses led to the appearance of evoked potentials (EP) with a latent period of 6–12 msec in the occipital, temporal, parietal, and cingular areas of the neocortex. The amplitude of the positive response component was 500 µV, and its duration 25–50 msec. The negative component was not always discernible. When rhythmic stimulation was used, these EPs followed stimulation frequencies not exceeding 20 per sec. Stimulation of the medial parts of the entorhinal area with a frequency of one to three per sec was accompanied by recruitment of the EP in the occipital and temporal neocortex areas. Nembutal depressed the amplitude of the neocortex EP appearing in response to stimulation of the entorhinal cortex. With the aid of double stimulation it could be established that, after conditioning stimulation of the entorhinal area, the positive component of the primary response (PR) evoked by stimulation of the contralateral sciatic nerve in the projection zone of the somatosensory cortex is strengthened during the first 50 msec, and subsequently after 80–120 msec. In these cases, the negative component was depressed. These findings are discussed with a view to the influence of limbic structures on the neocortex.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 1, pp. 73–78, January–February, 1970.     

Effect of transection of afferent pathways on evoked potentials,theta rhythm,and neuronal activity in the hippocampus

The effect of destruction of septo-hippocampal and subiculo-hippocampal connections on the electrical activity in the hippocampus was studied in tubocurarized rabbits. The theta rhythm and the so-called "intracellular theta rhythm" [7] were found to disappear on destruction of the septo-hippocampal connections, other consequences being the loss of neuronal capacity for responding to sciatic nerve stimulation, loss of neuronal inhibitory responses, and total or partial suppression of the negative phase of the evoked potential (EP). Destruction of subiculo-hippocampal connections entails a decrease in amplitude of the theta rhythm recorded from the hippocampal surface, retention of the "intracellular theta rhythm," and a slight decrease in amplitude of both EP phases. The number of neurons failing to respond to sciatic stimulation is increased; the character of cellular response remains unchanged. It is hypothesized that hyperpolarization of hippocampal pyramidal neurons may be responsible for generating the negative phase of the EP, and that this phase and the hippocampal theta rhythm may be of a common origin.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 4, pp. 439–444, July–August, 1970.     

Laminar analysis of hippocampal potentials evoked by peripheral stimulation

From areas SA₁ and SA₂ of the dorsal hippocampus of unanesthetized rabbits immobilized with d-tubocurarine a laminar analysis was made of evoked potentials (EP) in response to stimulation of the sciatic nerves. Inversion of the initial surface-positive phase of the EP was observed at the level of the pyramidal layer. The subsequent surface-negative phase reached a maximum value in the layer of basal dendrites of the pyramidal cells. The initial portion was inverted somewhat above the pyramidal layer and reached its maximum value approximately at the boundary of the pyramidal and radial layers. The change in sign of the remaining portion of this component occurred 0.3–0.4 mm deeper than the pyramidal layer. It is suggested that both components of the EP picked up from the hippocampal surface are due to an excitatory postsynaptic potential (EPSP) at the apical (positive phase), and basal (negative phase) dendrites. The positivity in the region of the pyramidal somata appears to be an extracellular reflection of a composite postsynaptic potential (IPSP) generated in this region.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 4. pp. 434–438, July–August, 1970.     

Neuronal firing in guinea pig neocortical slices surviving in vitro during adenosine-induced blockade of synaptic transmission

Background firing activity was recorded in guinea pig neocortical slices maintained using extracellular techniques. Between 30 and 40% of neurons continued to generate action potentials, although at a reduced rate, when synaptic disruption had been induced by adenosine or adenosine 5-monophosphate action. These cells were classed as endogenously active. No connection could be shown between neuronal firing pattern and capacity for autonomous generation of action potentials. The remaining neurons tested remained inactive after synaptic disruption, but regained their capacity for spontaneous firing following washout. The activity of these cells was classified as exogenous (or the result of synaptic excitation induced by other neurons in the same slice). The majority of cells with a highly regular discharge pattern initially stopped discharging during synaptic blockade and resumed their activity following washout. This would suggest that a miniature excitatory circuit with 30–140 msec cycles operates in these slices.Institute of Biological Physics, Academy of Sciences of the USSR, Pushchino. Translated from Neirofiziologiya, Vol. 19, No. 6, pp. 816–824, November–December, 1987.     

Analysis of long-latency components of field potentials evoked by stimulation of the cerebral cortex and nerves in the cerebellar paramedian lobule

Field potentials evoked in the graunular layer of the cerebellar paramedian lobule of unanesthetized cats in response to stimulation of the sensomotor cortex and limb nerves contained slow negative waves, appearing after a long latent period, which were generated by granule cells. In the case of nerve stimulation this component was recorded both inside and outside the projection zone of the corresponding limb. Cortical stimulation by single stimuli or series of stimuli not more than 1.8–2.5 times above threshold strength led to the appearance of evoked potentials only inside the corresponding projection zone. The long-latency component of field potentials evoked by cerebral stimulation followed high frequencies of repetitive stimulation and was less sensitive to the action of barbital anesthesia than the analogous component of potentials evoked by nerve stimulation. In the case of combined cerebral and nerve stimulation the long-latency components underwent summation. It is concluded that mossy fibers of slowly-conducting spino- and cerebrocerebellar tracts innervate different granule cells in the cerebellar cortex.Institute of Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 14, No. 4, pp. 379–385, July–August, 1982.     

Spatiotemporal characteristics of evoked potentials in the posterior ventrolateral nucleus of the cat thalamus

Evoked potentials (EPs) in the posterior ventrolateral nucleus (VPL) of the thalamus in response to electrical stimulation of the skin of the contralateral forelimb were studied. It was shown that they are formed by superposition of several electrical dipoles, one of which lies with two poles in VPL while the rest have one pole in VPL and the other in more caudal zones of the somatosensory system. The first phase of the EP in VPL consists of two components with different amplitude—frequency characteristics. The lability of the second component is low and it disappears if the frequency of stimulation rises above 20 Hz. The focus of maximal activity during contralateral stimulation of the skin of the forelimb occupies the same topographical position in VPL during development of the positive and negative phases of the EP. The steepness of rise of the ascending phase and the amplitude of the positive phase of the EP are greatest at the focus of maximal activity; in other regions the lines of equal steepness need not necessarily coincide with the equipotential lines; differences are especially noticeable in the sagittal plane.Institute of the Brain, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 4, No. 4, pp. 439–446, July–August, 1972.     

Electrophysiological indices of degree of functional integration between gratified neural tissue and the host brain

Neocortex from 17–18-day rat embryos was transplanted into the barrel-field cortical area in adult rats. Neuronal response to deflecting the vibrissae was tested in the graft 3–8 months afterwards. Nine out of 11 grafts showed a response to sensory stimulation. Irregular and asynchronous discharges predominated in neuronal background firing activity in these grafts. Generalized slow-wave activity had much in common with that occurring on the diametrically opposite site on the intact host brain cortex. Hypersynchronous volleys were detected in neurons of unresponsive grafts. A predominance of waves within the delta range while other rhythms remained only faint, together with epileptiform sharp spikes were seen in generalized activity. Histological treatment of responding grafts revealed close fusion between tissue and host brain. Non-responsive grafts were surronded by a thick glial scar.Institute of Biophysics, Academy of Sciences of the USSR, Pushchino-on-Oka. Institute of Neurobiology and Brain Research, E. German Academy of Sciences, Magdeburg, E. Germany. Translated from Neirofiziologiya, Vol. 19, No. 4, pp. 498–504, July–August, 1987.     

Spatial configuration of rabbit superior collicular evoked response to punctiform afferent stimulation

Evoked potentials arising in the rabbit superior colliculus in response to punctiform stimulation of the receptive field were studied. This response has only negative polarity at the focus of maximal activity and does not exhibit reversal of the potential which is a characteristic feature of the response to diffuse stimulation. The evoked potential was recorded at depths of between 0.1 and 0.9–1.0 mm from the collicular surface, corresponding to the stratum griseum superficiale. The response disappeared when the stimulating spot was shifted through 4–6° away from the optical position. It is suggested that evoked potentials to punctiform stimulation can give more complete information on the location of different synapses.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 11, No. 5, pp. 441–450, September–October, 1979.     

Effect of antibodies to glial glycolipid antigens on miniature endplate potentials

No more than 4% of the miniature endplate potentials (MEPP) recorded in preparations of frog cutaneous pectoral muscle kept in Ringers solution or treated with nonimmune rabbit serum were atypical, having a normal amplitude but abnormally retarded (almost doubled) time course. The percentage of atypical MEPP more than tripled after processing preparations with galactocerebroside (GalC) antiserum. Armine-induced blockade of synaptic acetylcholinesterase increased the rate of "giant" MEPP occurrence (but not of those with normal amplitude plus protracted time course). Mechanisms possibly underlying the increased percentage of atypical MEPP during GalC action on Schwann cells are discussed.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 21, No. 2, pp. 217–223, March–April, 1989.     

Analysis of the organization of afferent inputs in the projection cortex

Fast fluctuations in the evoked potentials (EP) at a local point of the projection cortex following stimulation of different pathways may reflect the activity of pyramidal neurons of different cortical layers. Analysis shows that the afferent and interarea projections to the somatic sensory cortex terminate on different neurons which can be regarded as relay neurons for a given pathway. Each group of neurons has its own system of inhibition for selective control of impulses coming along this pathway at the cortical level.Institute of Normal and Pathological Physiology, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 2, No. 4, pp. 368–372, July–August, 1970.     

Synaptic plasticity at archicortical and neocortical levels

Research carried out by the author and his collaborators, devoted to analysis of the properties and neurophysiological mechanisms of long-term (for several hours) potentiation, is surveyed. Long-term potentiation of focal potentials and unitary responses of strictly hippocampal structures (areas CA₁ and CA₃) in the unanesthetized rabbit is described. Enhancement of excitatory (EPSPs) and inhibitory (IPSPs) postsynaptic potentials was found after tetanization. No corresponding changes of sensitivity to acetylcholine or acetylcholinesterase activity were found by microiontophoretic and histochemical methods during long-term potentiation. Statistical analysis of EPSPs evoked by microstimulation, based on the quantal hypothesis of synaptic transmission, showed an increase in the number of quanta of transmitter release during potentiation. Long-term potentiation of focal potentials during stimulation of the subcortical white matter in surviving neocortical slices and also long-term potentiation of focal and unitary responses of the sensomotor cortex of the unanesthetized rabbit are described. Potentiation of the "indirect" component of the global response of the pyramidal tract was found. The data suggest the presence of long-term potentiation of monosynaptic neocortical responses. It is concluded that the main mechanism of both hippocampal and neocortical long-term potentiation is increased efficiency of excitatory synapses. It is postulated that synapses modified in this way are used in the formation of memory traces.Brain Institute, All-Union Mental Health Research Center, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 16, No. 5, pp. 651–665, September–October, 1984.     

Glial origin of the slow negative potential of the direct cortical response: Microelectrode study and mathematical analysis

The slow negative potential of the direct cortical response is similar in its shape, time course, and relationship to repetitive stimulation to depolarization of cortical glial cells but differs from the IPSP of cortical neurons. According to the results of digital spectral (frequency) analysis, the basis of the slow negative potential is the glial component formed by summation of components which coincide with glial depolarization processes with an accuracy determined by a constant factor. The much smaller component (as regards relative contribution) is the indirect result of the development of an IPSP in the neurons.I. S. Beritashvili Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 14, No. 1, pp. 76–84, January–February, 1982.     

Evoked potentials and unit activity in the cat somatosensory cortex

Investigation of unit activity of the cat somatosensory cortex has shown that the principal role in the genesis of the primary response, the response to stimulation of the thalamic relay nucleus, the callosal response, and certain other forms of evoked potentials (EPs) of the somatosensory cortex is played by neurons not usually responding by spike generation during EP development. The EPs reflect what the cortical neurons received from the afferent volley, and the level of their polarization, but they are not a reliable indicator of fast nervous processes in the cerebral cortex. The EPs reflect postsynaptic potentials (PSPs) of neurons not directly participating actively in the analysis of information reaching the cortex.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No, 4, pp. 360–367, July–August, 1970.     

Action of strychine on evoked potentials and postsynaptic responses of cat sensomotor cortical neurons

Acute experiments on immobilized cats lightly anesthetized with pentobarbital showed that application of strychine to the cortical surface inhibits slow negative potentials arising during direct and primary responses of the sensomotor cortex and corresponding IPSPs in pyramidal neurons. Iontophoretic applications of strychine blocks predominantly the early component of the IPSP, during which the input resistance under normal conditions is significantly less than during the late component of the IPSP, indicating that these components differ in their genesis. It is concluded that individual components of cortical evoked potentials have a common genesis, and that the slow negative potential is the dipole reflection of the IPSP in pyramidal neurons; the early component of the IPSP, moreover, is generated as a result of activation of axo-somatic inhibitory synapses, whereas the late component is generated as a result of activation of axo-dendritic synapses. The mediators in these inhibitory synapses may be different.I. S. Beritashvili Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 16, No. 4, pp. 480–487, July–August, 1984.     

Suppression of associative long-term potentiation in the hippocampus after tetanization of reinforcing afferents

Field potentials (FP) induced in area C1 by gentle orthodromic stimulation were recorded in murine hippocampal slices and associative long-term potentiation (ALTP) produced by C2 tetanization associated with intensive tetanization of another group of fibers (C1) was investigated. A comparison was made between the effects of additional C1 tetanization produced at 50–300 msec before and after combined tetanization of both afferents. Where these intervals measured 50–200 msec, preliminary tetanization of C1 suppressed ALTP (rise in FP amplitude: 10.4±5.2%) in comparison with the regimen whereby additional C1 tetanization came later (giving a rise of 32.4±5.3%); no significant difference was noted at an interval of 300 msec. The three possible reasons for ALTP suppression are discussed, namely: inactivation of "fast" calcium channels, post-activation hyperpolarization of postsynaptic neurons, and synaptic inhibition. The ALTP suppression mechanism is thought to resemble that underlying the relative inefficacy of "reversible" combinations in the shaping of behavioral conditioned reflexes.Institute for Brain Research, Academy of Medical Sciences of the USSR, Moscow. Institute of Chemical Physics, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 21, No. 5, pp. 636–643, September–October, 1989.     

Posttetanic changes in hippocampal minimal evoked potentials

Changes in the EEG induced by a single spike were recorded in the hippocampus of an unanesthetized rabbit. Summation of focal electrical activity synchronous with spontaneous single unit discharges at the symmetrical point of contralateral hemisphere revealed no stable potentials which could reflect these changes. In two cases discharges identified as activity of Shaffer's collaterals were recorded in area CA₁. Summation of post-spike changes in evoked activity recorded by the same microelectrode showed stable negative waves with an amplitute of 40–60 µV, which could have been evoked by single spikes. The curve of amplitude of the averaged evoked potentials versus near-threshold current strength stimulating the intrahippocampal pathways was not smooth in most experiments but stepwise in character. It is suggested that the minimal evoked potential corresponding to the first step (amplitude 40–80 µV) reflects a response to stimulation of one fiber. After above-threshold tetanization prolonged posttetanic potentiation of the minimal evoked potentials did not arise in CA₁ in response to stimulation of Shaffer's collaterals. Minimal evoked potentials recorded in area CA₃ in response to stimulation of the dentate fascia showed clear potentiation. The results are in agreement with the hypothesis of the synaptic localization of the mechanisms responsible for prolonged posttetanic potentiation.Brain Institute, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 9, No. 2, pp. 124–134, March–April, 1977.     

Time relations between the evoked potential and activity of hippocampal neurons

The time relations between the evoked potential (EP) and neuronal activity of the dorsal hippocampus in response to sciatic nerve stimulation were investigated in experiments conducted on rabbits paralyzed by tubocurarine. Two groups of neurons were distinguished on the basis of the type of their reaction to sciatic stimulation. Inhibition of background spike activity was found in the neurons of the first group (70.9%); in 37% of them inhibition was preceded by excitation in the form of a spike discharge or excitatory postsynaptic potential (EPSP) which coincided in time with the positive phase of the EP. During inhibition of spike activity the hyperpolarization potential was recorded intracellularly in a number of neurons, the latent period of which coincided with the latent period of the negative phase of the EP. Neurons of the second group (20%) were characterized by protracted excitation of spike activity, and the start of their excitation coincided with the start of the negative phase of the EP and hyperpolarization potential of the neurons of the first group. Different sensitivity of the two groups of neurons was noted. It is concluded that the EPSP of the pyramidal neurons of the hippocampus participates in generation of the positive phase of the EP, and the hyperpolarization potentials of these neurons participate in the generation of its negative phase. The possibility is not precluded that hippocampal neurons closer to the surface participate in the development of the negative phase of the EP.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 1, No. 3, pp. 285–292, November–December, 1969.     

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.     

The effect of the lateral reticular nucleus of the medulla oblongata on the cerebellar cortex

It was established as a result of a laminar analysis of evoked potentials (EP) in the paramedian lobe of the cerebellum of unanesthetized cats that in response to stimulation of the lateral reticular nucleus (LRN) excitation of granulosa cells develops which is not accompanied by excitation of Purkinje cells. Destruction of the lateral nucleus leads to a considerable decrease in the "diffuse" component of the EP which develop during stimulation of the somatic nerves. The results obtained show that the afferent pathway whose fibers innervate granulosa cells of a special type, which have been previously described, passes through the LRN. These cells have extensive receptive fields and do not directly affect the activity of Purkinje cells. A hypothesis is proposed on the basis of the results obtained and data in the literature that the extensive receptive fields of the LRN cells and of the granulosa cells innervated by them are due to interneuronal connections within the lateral nucleus.Institute of Problems of Information Transmission, Academy of Sciences of the USSR, Moscow. Laboratory on the Use of Mathematical Methods in Biology, Moscow University. Translated from Neirofiziologiya, Vol. 2, No. 6, pp. 581–586, November–December, 1970.     

Analysis of long cortical evoked potentials

Evoked potentials arising in the motor cortex in response to its direct stimulation (dendritic and slow negative potentials), to stimulation of the ventrolateral (primary response) and intralaminar (nonspecific response) thalamic nuclei, and to stimulation of the pyramidal tracts (antidromic response), and also postsynaptic responses of neurons corresponding to them were studied in acute experiments on curarized cats. Evoked potentials arising in response to direct cortical stimulation and also to stimulation of the specific and nonspecific thalamic nuclei and pyramidal tracts were recorded from the same point of the motor cortex, and the corresponding intracellular responses were recorded from the same neuron. Slow negative potentials arising under these conditions of stimulation and the IPSPs corresponding to them were shown to have an identical time course. The results show that slow negative potentials are a reflection of hyperpolarization of pyramidal neurons. It is suggested that the individual components of responses evoked by direct stimulation of the cortex and thalamic nuclei have a common genesis.I. S. Beritashvili Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 14, No. 2, pp. 115–121, March–April, 1982.