Dynamics of brain rhythmic and evoked potentials

In this study, the simultaneously recorded and selectively averaged evoked potentials in some of the structures of the auditory pathway (acoustical cortex, medial geniculate nucleus, inferior colliculus), in the reticular formation and the hippocampus of the awake cat as well as the simultaneous amplitude frequency characteristics of these structures are given. The power spectral density functions computed from the simultaneously recorded spontaneous activity of these structures are also presented. Using these results, the following analyses are accomplished: (1) determination of the dynamics of potentials simultaneously obtained from various brain structures, in order to evaluate the common features of their system characteristics; (2) determination of the relationship (or interactions) between rhythmic activity and evoked potentials of the brain, and (3) elaboration of a working hypothesis for the dynamics of potentials of the brain. Some suggestions and comments are also made for investigators working toward theories or dynamic models of signal transmission in the brain.     

Effect of local cooling of the cortex on evoked potentials in the reticular formation in response to somatosensory stimulation

Potentials evoked in nuclei of the reticular formation by electrodermal stimulation of the limbs were investigated in acute experiments on unanesthetized, immobilized rats during cooling of the somatosensory cortex in the area of representation of one forelimb. Evoked potentials in the reticular formation were found to depend on the degree of cold inhibition of the cortical primary response to the same stimulation. The peak time of the main negative wave increased from 40–50 to 60–80 msec with a simultaneous decrease in its amplitude or its total disappearance in the case of deep cooling of the cortex. Cooling of the cortex had a similar although weaker effect on the earlier wave of the evoked potential with a peak time of 14 msec, recorded in the ventral reticular nucleus. In parallel recordings of potentials evoked by stimulation of other limbs they remained unchanged at these same points of the reticular formation or were reduced in amplitude while preserving the same temporal parameters. Cooling of the cortex thus selectively delays the development and reduces the amplitude of the response to stimulation of the limb in whose area of representation transformation of the afferent signal into a corticofugal volley is blocked. Consequently the normal development of both late and early components of the potential evoked in the reticular formation by somatic stimulation requires an additional volley, descending from the cortex, and formed as a result of transformation of the same afferent signal in the corresponding point of the somatosensory cortex.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 13, No. 1, pp. 32–38, January–February, 1981.     

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.     

Dynamics of brain rhythmic and evoked potentials

In this study, the simultaneously recorded and selectively averaged evoked potentials in some of the structures of the auditory pathway (acoustical cortex, medial geniculate nucleus, inferior colliculus), in the reticular formation and the hippocampus of the cat during sleep as well as the simultaneous amplitude frequency characteristics of these structures are given. The power spectral density functions computed from the simultaneously recorded spontaneous activities of these structures are also presented. Using these results, the following analyses are accomplished:(1) determination of the dynamics of potentials simultaneously obtained from various structures, in order to evaluate the common features of their system characteristics; (2) determination of the relationship (or interactions) between rhythmic activity and evoked potentials of the brain.     

Effect of injury to the lateral reticular nucleus and nuclei of the inferior olive on electrical responses of the cerebellar cortex evoked by vagus nerve stimulation in cats

The role of the lateral reticular nucleus and nuclei of the inferior olive in the formation of cerebellar cortical evoked potentials in response to vagus nerve stimulation was determined in experiments on 28 cats anesthetized with chloralose and pentobarbital. After electrolytic destruction of the lateral reticular nucleus, in response to vagus nerve stimulation, especially ipsilateral, lengthening of the latent period and a decrease in amplitude of evoked potentials were observed; after bilateral destruction of this nucleus, evoked potentials could be completely suppressed. It is concluded that the lateral reticular nucleus relays interoceptive impulses in the vagus nerve system on to the cerebellar cortex. Additional evidence was given by the appearance of spike responses of Purkinje cells, in the form of mainly simple discharges, to stimulation of the vagus nerve. After destruction of the nuclei of the inferior olive, the latent period and the number of components of evoked potentials in response to vagus nerve stimulation remained unchanged but their amplitude was reduced. The role of the nuclei of the inferior olive as a regulator of the intensity of the flow of interoceptive impulses to the cerebellum is discussed.N. I. Pirogov Medical Institute, Vinnitsa. Translated from Neirofiziologiya, Vol. 9, No. 3, pp. 290–299, May–June, 1977.     

Human visual evoked potentials to bars and cross-like figures

Temporal and amplitude characteristics of averaged visual evoked potentials to presentation of lines, corners and crosses with different orientation recorded in 19 adult healthy subjects were compared in 34 derivations. In all subjects, the latencies of P1, N2, and P3 were shorter while their amplitudes were higher for crosses than for lines. The effect of lengthening of P1 peak latency from occipital to temporal cortex was mostly pronounced for EPs to a bar, whereas as increase in the P1 amplitude was most evident for a cross-like figure. Correlation of these data with: i) greater magnitude and shorter latencies of responses to crosses vs. bars in a half of cat striate neurons, ii) sensitivity of cells in monkey inferior temporal cortex to star-like figures, and iii) relatively better human recognition of figures with comers than with lines, as well as significance of the effects obtained for detection of image features in different areas of the human visual cortex is discussed.     

Variability of cortical evoked potentials to tooth pulp stimulation in unrestrained rats

In unrestrained adult rats evoked potentials were recorded by implanted electrodes in the somatosensory cortex in response to electrical stimulation of the pulp of an upper incisor. The spontaneous EEG, motor activity of the animal, and its respiratory movements were recorded simultaneously. Significant differences were observed in the configuration of the potentials and mean amplitude of the primary complex (P₁+N₁) during states of slow sleep, drowsiness, relaxed wakefulness, grooming, and investigative behavior; the amplitude of the primary complex during marked motor activity was reduced by more than an order of magnitude compared with that observed in a state of motor rest.In a state of relaxed wakefulness negative correlation was recorded between the amplitude of evoked potentials and momentary values of the respiration rate, weaker during periods of intensive motor activity. Meanwhile no direct parallel was observed between changes in potentials and respiration rate over the whole range of behavioral states studied: Depression of potentials was maximal during grooming whereas the respiration rate was maximal during investigative behavior.Paul Flechsig Institute for Brain Research, Karl Marx University, Leipzig, East Germany. Translated from Neirofiziologiya, Vol. 17, No. 1, pp. 27–35, January–February, 1985.     

Cerebral cortical respiratory-related evoked potentials elicited by inspiratory occlusion in lambs.

Respiratory-related evoked potentials (RREP) elicited by inspiratory mechanical loads have been recorded in humans. Early RREP peaks were hypothesized to be generated by activation of neurons in the somatosensory cortex. An animal model was developed to test this hypothesis in chronically instrumented, awake, spontaneously breathing lambs. Electrocorticogram (ECoG) was recorded bilaterally with ball electrodes on the dural surface over the somatosensory region. Inspiratory occlusions were presented through a face mask or endotracheal tube as interruptions of inspiration. Occlusion-elicited evoked potentials were obtained by computer-signal averaging the ECoG activity. A short-latency positive peak was observed bilaterally in the averaged occlusion-elicited evoked potentials in all animals breathing with the facemask and 5 of 8 lambs with the endotracheal tube. Postmortem identification of the electrode location demonstrated that the ECoG was recorded in the caudal-lateral portion of the somatosensory cortex. These results demonstrate that inspiratory occlusion elicits an evoked potential in the somatosensory cortical region of awake, spontaneously breathing lambs. The lamb cortical RREP is similar to human RREP.     

Distribution of evoked potentials on stimulation of the human pulvinar.

The distribution of the evoked cortical potentials recorded during stereotactic pulvinectomy is analyzed. The evoked cortical potential shows maximal amplitude in the precentral area, with decreasing amplitude in the parietal and anterior temporal area, and minimal amplitude in the occipital area. The pulvinar has been histologically considered to have dense connections with the parietal lobe and no connection with the frontal lobe. However, our results suggest that the pulvinar has a dense functional connection with the frontal cortex, through which the pulvinar plays a role in motor function.     

Midlatency auditory evoked responses: Differential effects of sleep in the cat

Middle latency responses (MLRs) in the 10–100 msec latency range, evoked by click stimuli, were studied in 8 adult cats during sleep-wakefulness to determine whether such changes in state were reflected by any MLR component. In particular, we wanted to determine whether the 20–22 msec positivity recorded at the vertex, ‘wave A,’ shown in previous studies to reflect a generator substrate within the ascending reticular formation, was tightly linked to changes in sleep-wakefulness, as reported for single neurons in the ascending reticular activating system. Evoked potentials were collected in 100 trial averages during continuous presentation of 1/sec clicks during initial awake recordings and thereafter during all-night sleep sessions. Continuously recorded EEG, EOG and EMG were scored for wakefulness, slow wave sleep (SWS), and rapid eye movement (REM) sleep during each evoked potential epoch. Recordings were obtained from electrodes implanted at the vertex and overlying the primary auditory cortex referenced to frontal sinus or to neck. In agreement with others, components of the auditory brain-stem response and the 12 msec primary cortical response showed no change in amplitude from wakefulness to either SWS or REM. Only wave A, among the components evaluated in the 1–100 msec range, decreased and disappeared during SWS and dramatically reappeared during REM to an amplitude equal to that during wakefulness. These data lend particular support to a functional relation between wave A and the ascending reticular activating system and suggest that this potential may provide a unique and dynamic probe of tonic brain activity. Moreover, this animal model provides a hypothetical basis for expecting a similar surface recorded potential in the human, a potential which has consequently been discovered.     

Neuronal generators of the visual evoked potentials: intracerebral recording in awake humans

Flash and pattern reversal visual evoked potentials were recorded in awake patients undergoing stereotactic procedures for severe dyskinetic disorders resistant to medical treatment. The nucleus ventralis lateralis thalami was reached via an occipital approach. VEPs were recorded on the scalp at the entracce of the intracerebral electrode, and serially from sites at different depths. A polarity reversal of the surface recorded wave form took place as the intracerebral electrode was advanced beneath the surface cortical layers. As concerns F-VEPs, most of the scalp activity mirrored the potentials recorded down to the depth of 70-65 mm from the thalamus. The largest amplitude of intracerebral F-VEPs was obtained from recording sites at 50–70 mm from the thalamus, i.e., in the depth of the calcarine fissure. A negative wave, peaking around 47–50 msec, became evident in recording sites at 30–40 mm from the thalamus but vanished as the electrode was advanced farther. In only one patient could we record a small negative wave, peaking at 33 msec, in the vicinity of the corpus geniculatum externum. Furthermore, the oscillatory activity recorded from the scalp appeared to be generated in the cortical layers. PR-VEPs also underwent polarity reversal as the electrode traversed the cortex. PR-VEPs disappeared more superficially than F-VEPs. No PR-evoked activity could be recorded in the vicinity of the corpus geniculatum externum.We conclude that slow and fast components of VEPs recorded from the scalp are entirely generated in cortical layers.     

Study of the evoked potentials to photic stimulation in man after deafferentiation of the visual cortex]

A study was made of evoked potentials (EP) in response to light stimulus in the occipital areas of the cortex in 28 patients with homonimous hemianopsia caused by focal affection of the optic system. EP asymmetry in the hemispheres was recorded only in case of complete homonimous hemianopsia. Reduction of response at the side of contralateral hemoanopsia was observed in affection of the optic tracts at the level of the optic cord and in case of spread of the pathological focus to the occipital lobe. In half of the cases of homonimous hemianopsia caused by affection of the optic radiation the responses were greater by amplitude at the side of the focus or were recorded only in the deafferentated occipital area.     

Characteristics of interhemispheric relationships in the absence or presence of a skill

A comparative analysis of the time and amplitude characteristics of the negative N200 and positive P300 components of visual evoked potentials recorded at symmetric points of the frontal, parietal, temporal, and occipital areas of the right and left hemispheres of the cerebral cortex has been performed in subjects with or without the skill of operating a computer. Subjects inexperienced in an operator’s work exhibited an interhemispheric difference in the time and amplitude characteristics of the studied components. In subjects that had the skill of operating a computer, the interhemispheric difference was little, which suggests that the cortex plays only a small role in the cerebral control of this activity.     

Commissural channels of visual information transmission

Experiments were made in adult cats with different transections of the classical and commissural visual tracts to study evoked potentials and unit neuronal activity in response to visual stimulation. The commissural channels of the telencephalon, diencephalon and mesencephalon were demonstrated to be highly effective in visual information conduction to the visual cortex. Complete transection of the classical and commissural tracts with the exception of the commissural tract of the third ventricle fundus and midbrain reticular formation resulted in the disappearance of evoked potentials in the visual cortex in response to light flashes, despite the fact that 8 of the 15 recorded neurons responded to photostimulation. Activation of such neurons was caused by visual information conduction via two possible commissural channels: midbrain reticular formation and subventricular interhemispheric commissures of the diencephalon.     

Electrophysiological study of the functional connections of the hypothalamus with the forebrain in the tortoise Emys orbicularis

During stimulation of the posterior hypothalamus, the evoked potentials with short latent periods, high amplitude and poor exhaustion by rhythmic stimulation were recorded in the hippocampal cortex. In the piriform cortex, the evoked potentials exhibited longer latent periods and complex configuration. Less readily the evoked potentials appeared in the neocortex, their latency being very large. During stimulation of the anterior hypothalamus, maximum activity was also localized in the hippocampal cortex. The data obtained indicate close connection between hypothalamic structures and the hippocampal cortex. The latter is presumably the main projectional area for the ascending afferentation from the hypothalamus.     

Comparative analysis of the role of the second and first somatosensory cortical areas in somatic responses of the medullary reticular formation in rats

Evoked potentials and unit activity in the medullary reticular formation were investigated in unanesthetized, curarized rats during cold blocking or after extirpation of the cortical representation of one of the stimulated limbs. Local cooling or extirpation in area SII, unlike blocking of area SI, leads to a small (up to 30%) decrease in amplitude and a very small change (up to 10 msec) in the temporal parameters of evoked potentials arising in the reticular formation in response to electrodermal stimulation of the contralateral limb, whose representation in the cortex was blocked. Predominance of corticofugal influences from SI over those from SII was discovered both in experiments with evoked potentials and during analysis of somatic spike responses of reticular formation neurons. Corticofugal control over activity of the medullary reticular formation in rats exerted by neuronal mechanisms of somatosensory areas SII and SI thus differs both qualitatively and quantitatively.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 15, No. 1, pp. 42–49, January–February, 1983.     

Enhanced Stimulus-Induced Gamma Activity in Humans during Propofol-Induced Sedation

Stimulus-induced gamma oscillations in the 30–80 Hz range have been implicated in a wide number of functions including visual processing, memory and attention. While occipital gamma-band oscillations can be pharmacologically modified in animal preparations, pharmacological modulation of stimulus-induced visual gamma oscillations has yet to be demonstrated in non-invasive human recordings. Here, in fifteen healthy humans volunteers, we probed the effects of the GABA_A agonist and sedative propofol on stimulus-related gamma activity recorded with magnetoencephalography, using a simple visual grating stimulus designed to elicit gamma oscillations in the primary visual cortex. During propofol sedation as compared to the normal awake state, a significant 60% increase in stimulus-induced gamma amplitude was seen together with a 94% enhancement of stimulus-induced alpha suppression and a simultaneous reduction in the amplitude of the pattern-onset evoked response. These data demonstrate, that propofol-induced sedation is accompanied by increased stimulus-induced gamma activity providing a potential window into mechanisms of gamma-oscillation generation in humans.     

Effects of morphine on visual evoked responses recorded in five brain sites.

The effects of morphine on averaged evoked responses to visual stimulation were examined in specific brain structures relevant to pain, analgesia, tolerance and motor disturbances. Permanent electrodes (60 μ in diameter) were implanted stereotaxically in the central gray, mesencephalic reticular formation, caudate nucleus, parafasicular-centromedian complex and the lateral geniculate body as a control site. Visual evoked responses were obtained in unanesthetized, unrestrained rats prior to and following the administration of morphine in successive doses of 1, 5, 10 and 30 mg/kg and 1 mg/kg of naloxone (a morphine antagonist). The parafasicular-centromedian complex and the reticular formation exibited a progressive increase in response amplitude to increased dose of morphine. These effects were reversed by naloxone. In this study the parafasicular-centromedian complex was found to be the most sensitive structure to morphine, displaying the largest changes in response amplitude as a result of morphine administration.     

Ontogenetic patterns of frequency-power spectra elicited by flickering light in children: a comparative study of evoked and background EEGs.

Frequency-power spectra of the EEG evoked by repeptitive photic stimulation and of the background EEG were studied during childhood in 43 awake subjects aged between 2 months and 14 years. EEG activity was recorded from the middle parieto-occipital region with the aid of a 1-channel analyzer Lysograf-Alvar, analysing 16 frequencies in the range from 2 to 28 c/sec. The responsiveness of the central nervous system to flickering light improved in the course of childhood in parallel with the significant decline of delta activity and with the prominent increase of alpha intensity in the resting EEG. The 4th month of life appeared to be a marked turning point in the development of evoked and background EEGs. From that age, the bioelectric power at the flash rate corresponding to photic "driving" began to increase together with the highest and optimal driving frequencies. The flash rate, at which evoked potentials changed into the "driven" rhythm, also shifted towards higher frequencies. Subsequently, the amount of energy in the resting EEG increased significantly within the theta, alpha and beta bands and, on the contrary, a prominent decline was observed in the delta range. Marked ontogenetic changes at this age closely coincided with the rapid development of exogenous fibres in the occipital cortex, including the thalamo-cortical conncetions, and fibres of the neuropil in cortical layer I, which might play an important role in the genesis of background and "driven" in the occipital region.     

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.