Cortical and subcortical evoked potentials and behavioral responses after destruction of the globus pallidus in CATS

Evoked potentials (EPs) to clicks were recorded in the ectosylvian and sigmoid gyri of the cortex, in the medial geniculate body, and in the median nucleus of the thalamus by means of permanently implanted electrodes in experiments on 14 cats. A motor component of the behavioral response (conditioned food reflex) developed in response to the same stimulus (clicks). In the first group of animals the region of destruction was confined to the globus pallidus. In these animals EPs to clicks appeared during the first days after the operation in all structures, although in some tests they were reduced. Meanwhile the behavioral response to clicks was absent and showed no signs of spontaneous recovery. It could be formed (retraining) later in the experiments. In the second group of animals the globus pallidus was not completely destroyed, but neighboring areas of the entopeduncular nucleus and internal capsule were coagulated. In these cats more marked changes in behavioral and electrophysiological indices were observed. EPs to clicks disappeared in some cases for 25–30 days, subsequently returning to normal. The role of the globus pallidus as a "non-specific" polymodal structure in the analysis of sensory information is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 2, pp. 141–149, March–April, 1972.     

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.     

Features of evoked potentials in the cerebellar cortex of rabbits

We investigated evoked responses of the cerebellar cortex of rabbits under Nembutal or chloralose anesthesia to stimulation of the sciatic, brachial, and vagus nerves. The parameters of evoked potentials (E Ps), together with features of their distribution throughout the cerebellar cortex, enabled us to divide them provisionally into three types. Evoked potentials of the first type have a latent period of 5–10 msec and a two-phase or more complex shape. Evoked potentials of the second type have a latent period of 10–23 msec and include from one to four components. Evoked potentials of the third type are discharges with long latent periods (20–50 msec) and consist of a series of slow sinusoidal oscillations. Appearance of an initial electronegative component is characteristic of EPs of the cerebellar cortex of rabbits, especially those of the second and third types. Evoked potentials of the first type are local.N. I. Pirogov Vinnitsa Medical Institute. Translated from Neirofiziologiya, Vol. 1, No. 1, pp. 73–80, July–August, 1969.     

Electrical activity of thalamocortical association systems in postnatal ontogeny of the cat

Electrical responses to somatic, photic, and acoustic stimulation in the sensomotor, parietal, temporal, and occipital regions of the cortex were studied in the nucleus lateralis posterior and nucleus ventralis lateralis of the thalamus by recording averaged evoked potentials in kittens (aged 3 to 41 days) anesthetized with pentobarbital. A definite order of maturation of afferent inputs into cortical association areas was demonstrated. The parietal cortex was shown to become polysensory before the sensorimotor cortex. It is suggested that the nucleus lateralis posterior is the main thalamic nucleus responsible for conduction of visual information to the cortex in kittens during the first month of life. Incorporation of this nucleus into the system conducting somatic impulsation to the sensorimotor cortex takes place by the age of 3 weeks.A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 14, No. 5, pp. 476–482, September–October, 1982.     

Changes in amplitude and temporal characteristics of evoked potentials in the sensomotor cortex after division of the spinocervical tracts in cats

Temporal and amplitude characteristics of evoked potentials of the sensomotor cortex in waking cats were studied during variation in the intensity of electrodermal stimulation. The results obtained in experiments on intact animals and on the same animals for several months after division of the spinocervical tracts at the cervical level were compared. After blocking of the inflow of afferent impulses along these tracts of the spinal cord, statistically significant changes in evoked potentials were observed, mainly in response to medium and strong stimulation. These changes were more clear in the motor and second somatosensory areas of the cortex. A decrease in sensitivity to pain also was found. During recovery of the motor functions, cutaneous sensation remained impaired and the amplitude characteristics of the evoked somatosensory activity were not restored. The results suggest that thinner fibers predominate among the primary afferent fibers of the spinocervical tract, and their projections are more widely represented in the second somatosensory and motor areas of the cortex.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 4, No. 5, pp. 516–523, September–October, 1972.     

Effect of peripheral stimulation on neocortical electrical activity in the chronic premesencephalic cat

Chronic experiments on 19 adult cats showed that on the second or third day after premesencephalic section through the brain stem preserving the lemniscal pathways, auditory, olfactory, and cutaneous stimulation evoke diffuse neocortical activation. Although photic stimulation evoked responses in the visual cortex, it could not desynchronize the neocortical activity. It is postulated that diffuse neocortical activation in cats with a complete block of the mesencephalic recticular formation in response to auditory, olfactory, and cutaneous stimulation arises through excitation of the posterior hypothalamus.Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 7, No. 5, pp. 493–499, September–October, 1975.     

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.     

Mechanisms of interaction between the parietal association and projection areas of the cat neocortex

Changes in evoked potentials in the first visual (VI), first somatic (SI), and parietal areas of the cortex during local cooling of each area were investigated under pentobarbital anesthesia. Two types of interaction were distinguished. Type I interaction was found in all areas in the early stages of local cooling and was reflected in a similar decrease in amplitude of evoked potentials in intact parts of the cortex. In the thalamic association nuclei — the pulvinar and posterolateral nucleus — somatic evoked potentials were unchanged but visual were transformed differently from those in the cortex. Type IIinteraction was found in the later stages of cooling and only between the association area and each of the projection areas. It was reflected in a greater change in amplitude of the evoked potentials and also in their configuration. In response to somatic stimulation in the early stage of type II interaction transformation of evoked potentials in the cortex took place sooner than in the nuclei; in the later stage it took place immediately after transformation of the "subcortical" evoked potentials. In response to photic stimulation transformations of cortical evoked potentials were always preceded by the corresponding transformations in the nuclei. It is suggested that type I interaction is formed by intercortical connections and type II by direct and subcortical relay connections. Differences in the role of the association area in interaction of types I and II when activated by stimuli of different modalities are discussed.Brain Institute, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 10, No. 6, pp. 573–581, November–December, 1978.     

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.     

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.     

Relevance of the callosal transfer in defining the peripheral reactivity of somesthetic cortical neurones

Experiments were performed in 13 chloralose-anaesthetized, curarized cat preparations (monitoring of rectal temperature, heart rate, expired pCO2 and EEG), in order to ascertain whether, and to what extent, the reactivity to ipsilateral skin shocks of the neurones of the anterior ectosylvian and anterior suprasylvian gyri (AEG and ASG, respectively) is dependent on the callosal output of the somatosensory areas of the contralateral hemisphere. Indeed, we knew from previous experiments that a high proportion of AEG and ASG neurones having bilateral peripheral receptive fields (PRFs) can be excited by direct stimulation of the contralateral homonymous areas, and that the callosal fibres originating in the latter carry somesthetic impulses related to contralateral PRFs. A preliminary analysis was carried out on the amplitude and latency relationships between the evoked potentials (EP) recorded simultaneously from the two hemispheres and from the corpus callosum (CC) following stimulation of the forepaw of one side. The results obtained showed good correlations between the onset and development of the EPs picked up from the hemisphere ipsilateral to the stimulated skin, on the one hand, and onset and development of the EPs recorded from the contralateral hemisphere and the corpus callosum, on the other. At a further stage of the experiments, the EPs elicited upon ipsilateral and contralateral skin shocks in the AEG-ASG area have been recorded and averaged before, during and after the reversible inactivation of callosal somesthetic transmission. This was achieved by applying polarizing currents (0.2-1 mA) to the rostral portion of the CC, adequacy and reversibility of this method having been tested by observing, respectively, suppression and prompt restoration of transcallosal EPs and of the asynaptic spiking produced by cortical cells when antidromically invaded from contralateral homotopic cortex. It was seen that during CC blockade the EPs elicited in the AEG-ASG areas did not show any change either in amplitude or time-course if brought about by contralateral peripheral stimulation, whereas those evoked by ipsilateral skin shocks exhibited significant reduction, which was related to the strength of CC polarization and to the reduction of transcallosal EPs. In control experiments similar effects were observed after ablation of somatosensory areas of the hemisphere which send off somesthetic callosal impulses, whereas strychninization of these areas caused effects opposite in sign, i.e., enhancement of the ipsilateral but not of the contralateral EPs in the areas of the untreated hemisphere. By testing the effects of CC polarization on single AEG-ASG neurones, it was observed that the responses of units linked only with contralateral PRFs (Group I; 7 units tested) were unaffected by callosal polarization. The discharges of neurones provided with wide and bilateral PRFs (Group II; 27 units tested) were not affected if elicited by contralateral PRF shocks but were deeply impaired (in 11 neurones out the 27) when provoked by ipsilateral PRF stimulation. The effect consisted chiefly of the disappearance of the first high peak of the PSTHs, and when recording intracellularly graded events, it was mirrored by a large decrease of the postsynaptic excitatory potentials elicited in Group II neurones by ipsilateral PRF shocks. A late scattered histographic component was identified in the PSTHs of such cells, which did not appear to be significantly altered during CC blockade. These effects were observed on the ipsilateral responses of 11 out of the 27 Group II neurones so tested whereas the ipsilateral PSTHs of the remaining 16 Group II neurones either did not undergo significant changes during the callosal blockade or escaped evaluation because of high spontaneous shifts of neural responsiveness. The results are discussed mainly with a view to the possible functional role of the specific somesthetic callosal fibres in defining ipsilateral reactivity for the wide-field cells of the AEG-ASG area.     

Brain activation by photic stimulation in normal cats and after transection of afferent pathways to the mesencephalic reticular formation

In chronic experiments on waking, unimmobilized cats with implanted electrodes, the EEG activation reaction evoked by short flashes of varied intensity was recorded in normal animals and after blocking excitatory influences of the visual sensory system on the brain-stem reticular formation by transection of the brachium colliculi superioris bilaterally. The intensity of the activation reaction in intact animals increased steadily with an increase in intensity of the photic stimuli. No clear dependence of response amplitude on stimulus intensity was observed in the cats with transection; the changes were largely random in character. It is suggested that these disturbances were due to interruption both of the ascending flow excitatory afferent impulses to the brain-stem reticular formation through the brachia and superior colliculi and of corticofugal volleys descending from the cortex to the mesencephalic reticular formation via the same brachia, and controlling the level of its activation.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 13, No. 5, pp. 500–505, September–October, 1981.     

Comparison of evoked potentials to tones and clicks recorded simulataneously by multiple electrodes from the auditory cortex in unanesthetized cats

Evoked potentials to tones and clicks were recorded simultaneously from seven points of the auditory cortex and one or two points of the somatosensory cortex in unanesthetized cats. Comparison of evoked potentials to tones of equal loudness in the 250–7000 Hz band showed no common pattern of cortical tonotopic distribution. However, an individual dependence of the components of the evoked potential on pitch and on localization of the recording point exists for each animal. With a change in stimulus intensity the absolute and relative values of these components of the evoked potential vary. The initial positive waves are the most variable; besides the two waves already known a third, intermediate wave, particulary sensitive to loudness, was discovered. The negative wave of the primary response increases proportionally to loudness. Evoked potentials to clicks are more uniform over the auditory cortex and more stable than those to tones. Responses appeared in the somatosensory cortex to loud stimuli, more regularly to clicks than to tones. It is concluded that the parameter of pitch is reflected in the cat cortex as a complex spatially-individual distribution of the amplitude and time parameters of the evoked potentials.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 7, No. 2, pp. 115–125, March–April, 1975.     

The effects of neck flexion on cerebral potentials evoked by visual,auditory and somatosensory stimuli and focal brain blood flow in related sensory cortices

Background

A flexed neck posture leads to non-specific activation of the brain. Sensory evoked cerebral potentials and focal brain blood flow have been used to evaluate the activation of the sensory cortex. We investigated the effects of a flexed neck posture on the cerebral potentials evoked by visual, auditory and somatosensory stimuli and focal brain blood flow in the related sensory cortices.

Methods

Twelve healthy young adults received right visual hemi-field, binaural auditory and left median nerve stimuli while sitting with the neck in a resting and flexed (20° flexion) position. Sensory evoked potentials were recorded from the right occipital region, Cz in accordance with the international 10–20 system, and 2 cm posterior from C4, during visual, auditory and somatosensory stimulations. The oxidative-hemoglobin concentration was measured in the respective sensory cortex using near-infrared spectroscopy.

Results

Latencies of the late component of all sensory evoked potentials significantly shortened, and the amplitude of auditory evoked potentials increased when the neck was in a flexed position. Oxidative-hemoglobin concentrations in the left and right visual cortices were higher during visual stimulation in the flexed neck position. The left visual cortex is responsible for receiving the visual information. In addition, oxidative-hemoglobin concentrations in the bilateral auditory cortex during auditory stimulation, and in the right somatosensory cortex during somatosensory stimulation, were higher in the flexed neck position.

Conclusions

Visual, auditory and somatosensory pathways were activated by neck flexion. The sensory cortices were selectively activated, reflecting the modalities in sensory projection to the cerebral cortex and inter-hemispheric connections.     

Tonic and phasic modifications of viscerosensory evoked potentials during sleep in cats

Modification of the viscerosensory evoked potentials (EPs) were studied during the sleep-wakefulness cycle of the rat. Electrical stimuli of various intensity were delivered either to the mucosal surface of a fistula of the small intestine or to the left splanchnic nerve during wakefulness (W), drowsiness (D), slow-wave-sleep (SWS), and paradoxical sleep (PS). The average EPs were recorded from the somatosensory (SI and SII) and associative (AS) areas of the cortex, the ventrobasal complex of the thalamus (VPL), the posterior hypothalamus (HPT) and the dorsal hippocampus (HPC). The amplitude of each component of the EPs in all explored structures were the largest in SWS and the smallest in W. A phasic increase in amplitude was observed in the EPs recorded immediately before the appearance of the spindles of SWS and during the REM episodes of PS. The peak latencies of the late components were the longest in SWS. These changes of the amplitudes and latencies were greater in the responses to weak stimulation than in EPs to strong ones. The possible synaptic events of the sleep-dependent control of viscerosensory activity are discussed.     

Spread of fast thermowaves through the cerebral cortex produced by visual stimulation

Thermowaves spreading through the cerebral cortex were observed and investigated during acute experiments on white rats using a new technique — that of thermoencephaloscopy (TES) through the intact skull. These waves were induced by regular visual stimulation (at intervals of 1.5–3 min) or by initiating one of the flashes and spread through the cortex along trajectories of five basic types; amplitude of thermowaves: 0.005–0.1°C; length: 10–15 mm; duration 1.2–11.4 sec; velocity: 1–33 mm/sec; extent of pathway: 2–56 mm. They appeared with a high degree of probability (of 0.92) during the interval between 15 sec before and 26 sec after the flash. Numbers of moving waves declined by the point of stimulus application and rose by 7–8 sec after the flash. Waves arose in 50% of cases in the contralateral visual cortex (areas 17 and 18a), spreading to the midline and crossing to the ipsilateral hemisphere (areas 17, 18a, and 7). Local waves moving along a circular trajectory were also discovered in the contralateral visual cortex. Several types of wave, differing in trajectory, also arose in the ipsilateral visual cortex. Mechanisms and possible significance of this effect are examined.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 21, No. 4, pp. 467–475, July–August, 1989.     

Formation of visual evoked potentials and development of visually guided behavior in the Ficedula hypoleuca nestling

Age dynamics of evoked potentials (EPs) of Wulst area of the dorsal hyperstriatum (functional analogue of mammalian visual cortex) was investigated in 2-7 days old nestlings. EPs in response to light stimuli, with the duration and intensity imitating natural alimentary signals for 5-7 days-old nestlings, were recorded parallel with the behaviour. It was found that the development of the visual system in precocious nestlings became completed in postembryonal ontogenesis. The most essential changes in the parameters of visual EPs occur between 2.5 and 4 days of nest life. By the end of the 4-th day the EPs parameters resemble those of the definite EPs. During the same period the visually directed form of the alimentary behaviour appears for the first time. Both the visual EPs and the visually directed alimentary behaviour first appear in the ontogenesis in response to the second component of two-fold luminosity change--the natural signal of alimentary behaviour.     

Motor Representations of Facial Muscles and Vibrissae in Cerebellar Cortex of the White Mouse Mus musculus

Facial motor responses (MR) to cerebellar cortex microstimulation (MS) were studied in scalped, hexenal-anesthetized white mice. Series of MS as 7 impulses with frequency 333 imp/s and current 10–50 µA evoked predominantly ipsilateral MR in vibrissa groups, upper lip, external ear, mandible, and eyeball. Facial motor representations (FMR) are expressed as numerous, small-volume areas located mainly in vermix and adjacent areas of cerebellar cortex. The number of facial tracks rises with increase of depth of microelectrode inserion. FMR in the white mouse cerebellar cortex are asymmetrical, with prevalence of facial zone in the left cerebellar part. It is suggested that there is a common relay link for cortico-, tectal-, and cerebellobulbar pathways of regulation of facial motor activity and that the role of cerebellum consist in modulation of motor impulses from cortico- and tectofacial systems, which provides a finer coordination of different muscle groups for the whole motor act realization.     

Compression and reflection of visually evoked cortical waves

Neuronal interactions between primary and secondary visual cortical areas are important for visual processing, but the spatiotemporal patterns of the interaction are not well understood. We used voltage-sensitive dye imaging to visualize neuronal activity in rat visual cortex and found visually evoked waves propagating from V1 to other visual areas. A primary wave originated in the monocular area of V1 and was "compressed" when propagating to V2. A reflected wave initiated after compression and propagated backward into V1. The compression occurred at the V1/V2 border, and local GABAA inhibition is important for the compression. The compression/reflection pattern provides a two-phase modulation: V1 is first depolarized by the primary wave, and then V1 and V2 are simultaneously depolarized by the reflected and primary waves, respectively. The compression/reflection pattern only occurred for evoked waves and not for spontaneous waves, suggesting that it is organized by an internal mechanism associated with visual processing.     

Postsynaptic potentials of facial motoneurons evoked by afferent and corticofugal impulses

Postsynaptic potentials of motoneurons in the facial nerve nucleus, evoked by stimulation of the cranial nerves (trigeminal, hypoglossal, facial) and of the sensomotor cortex were investigated in cats anesthetized with chloralose and pentobarbital. Two functionally opposite groups of motoneurons were found to exist in the facial nucleus. Stimulation of the afferent nerves and cortex evoked the appearance of EPSPs in the first of these groups and IPSPs in the second. The latency and duration of the PSPs indicate that afferent and corticofugal impulses reach the facial motoneurons along polysynaptic pathways. Interneurons on which wide convergence of influences travelling along afferent fibers and of the cortex, were found in the region of the facial nucleus. The possible neuronal pathways concerned with the transmission of afferent and corticofugal impulses to the facial motoneurons are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol.4, No.4, pp. 391–400, July–August, 1972.