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.     

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.     

A study of the time and frequency characteristics of the potentials evoked in the acoustical cortex

Evoked potentials in the auditory cortex of the cat are measured by applying auditory stimulations in the form of tone bursts of 700 Hz. Transient evoked potentials obtained in this way are transformed to the frequency domain using a Laplace Transform. The amplitude frequency characteristic obtained with this semi-empirical method depicts maxima of EEG-amplitude in frequency ranges of 10–13 Hz and 60–80 Hz. The correlation between the time course of evoked potentials and spontaneous activity of the brain and the efficiency of the method used are pointed out.     

Evoked potentials of the auditory cortex of the porpoise,Phocoena phocoena

Summary Evoked potential (EP) recordings in the auditory cortex of the porpoise,Phocoena phocoena, were used to obtain data characterizing the auditory perception of this dolphin. The frequency threshold curves showed that the lowest EP thresholds were within 120–130 kHz. An additional sensitivity peak was observed between 20 and 30 kHz. The minimal EP threshold to noise burst was 3·10^–4–10^/s-3 Pa. The threshold for response to modulations in sound intensity was below 0.5 dB and about 0.1% for frequency modulations. Special attention was paid to the dependence of the auditory cortex EP on the temporal parameters of the acoustic stimuli: sound burst duration, rise time, and repetition rate. The data indicate that the porpoise auditory cortex is adapted to detect ultrasonic, brief, fast rising, and closely spaced sounds like echolocating clicks.Abbreviation EP evoked potential     

Responses of neurons in the rat auditory and associative cortices to tonal stimuli

Activity of single neurons and mass evoked potentials (EP) were recorded from the auditory (area 41) and associative (area 39) cortices in acute experiments on rats anesthetized with urethane, nembutal, or chloralose; pure tones were used as acoustic stimuli. The EP appearing in response to a wide range of sound tones on the surface of the auditory and associative cortices were dissimilar in their latency and shape. For neurons exhibiting stable responses, the frequency-threshold curves (FTC) were plotted.Weak and variable responses of neurons were observed under slight urethane anesthesia. Nembutal anesthesia increased the responsiveness of neurons and the probability of appearing of late components in the responses. Chloralose anesthesia was characterized by extension of frequency range perceived by a neuron, while its sharpness of tuning remained unchanged. Under all types of anesthesia employed, the responses recorded from the associative cortex neurons had longer latencies than those recorded from the auditory cortex neurons. Neurons exhibiting the frequency selectivity were much less numerous in the associative cortex than in the auditory cortex. The former neurons were often characterized by intermittent FTC and they responded to a more extended frequency range. No clear tonotopic organization was found in the associative cortex.Neirofiziologiya/Neurophysiology, Vol. 25, No. 5, pp. 343–349, September–October, 1993.     

Interaction of influences from the auditory and somatosensory afferent systems on neurons of the primary auditory cortex

In experiments on anesthetized cats, 80 neurons of the primary auditory cortex (A1) were studied. Within the examined neuronal population, 66 cells (or 82.5%) were monosensory units, i.e., they responded only to acoustic stimulations (sound clicks and tones); 8 (10.1%) neurons responded to acoustic stimulation and electrocutaneous stimulation (ECS); the rest of the units (7.4%) were either trisensory (responded also to visual stimulation) or responded only to non-acoustic stimulations. In the A1 area, neurons responding to ECS with rather short latencies (15.6–17.0 msec) were found. ECS usually suppressed the impulse neuronal responses evoked by sound clicks. It is concluded that somatosensory afferent signals cause predominantly an inhibitory effect on transmission of an acoustic afferent volley to the auditory cortex at a subcortical level; however, rare cases of excitatory convergence of acoustic and somatosensory inputs toA1 neurons were observed.     

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.     

Sensory and cognitive evoked potentials in a case of congenital hydrocephalus

We studied auditory and visual evoked potentials in D.W., a patient with congenital stenosis of the cerebral aqueduct. Head CT scans revealed marked hydrocephalus with expanded ventricles filling more than 80% of the cranium and compressing brain tissue to less than 1 cm in thickness. Despite the striking neuroanatomical abnormalities, however, the patient functioned well in daily life and was attending a local community college at the time of testing.Evoked potentials provided evidence of preserved sensory processing at cortical levels. Pattern reversal visual evoked potentials had normal latencies and amplitudes. Brain-stem auditory evoked potentials (BAEPs) showed normal wave V latencies. Na and Pa components of middle-latency AEP had normal amplitudes and latencies at the vertex, although amplitudes at lateral electrodes were larger than at the midline.In contrast to the normal sensory responses, long-latency auditory evoked potentials to standard and target tones showed abnormal P3 components. Standard tones (probability 85%), evoked NN1 components with normal amplitudes (−3.7 μV) and latencies (103 msec), but also elicited large P3 components (17 μV, latency 305 msec) that were never observed following frequent stimuli in control subjects. Target stimuli (probability 15%) elicited P3s in D.W. and controls, but P3 amplitudes were enhanced in D.W. (to more than 40 μV) and the P3 showed an unusual, frontal distribution. The results are consistent with a subcortical sources of the P300. Moreover, they suggest that the substitution of controlled for automatic processes may help high-functioning hydrocephalics compensate for abnormalities in cerebral structure.     

Responses of units in the magnocellular part of the medial geniculate body to acoustic and somatosensory stimulation

Responses of 150 neurons in the magnocellular part of the medial geniculate body to clicks and to electrodermal stimulation of the contralateral forelimb were investigated in cats immobilized with myorelaxin. Of the total number of neurons 65% were bimodal, 16.6% responded only to clicks, and 15.4% only to electrodermal stimulation. The unitary responses were excitatory (spike potentials) and inhibitory (inhibition of spontaneous activity). Responses beginning with excitation occurred more frequently to stimulation by clicks than to electrodermal stimulation, whereas initial inhibition occurred more often to electrodermal stimulation. The latent period of the initial spike potentials in response to clicks and to electrodermal stimulation was 5–27 and 6–33 (mean 11.6 and 16.2) msec respectively. Positive correlation was found between the latent periods of spike potentials recorded in the same neurons in response to clicks and to electrodermal stimulation, and also to electrodermal stimulation and to stimulation of the dorsal funiculus of the spinal cord. It is concluded that the magnocellular division of the medial genicculate body is a transitional structure between the posterior ventral nucleus and the parvocellular division of the medial geniculate body, and that in addition, it is connected more closely with the auditory than with the somatosensory system. It is suggested that the somatosensory input into the magnocellular division of the medial geniculate body is formed mainly by fibers of the medial lemniscus.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 2, pp. 133–141, March–April, 1978.     

Mesencephalic chronic electrodes in pain patients. An electrophysiological study.

Early components of lemniscal potentials after contralateral median nerve or mechanical stimulus are due to lemniscal pathways, whereas later components, after 70 msec appearing bilaterally and at higher stimulus intensities probably express extralemniscal activity. Evoked potentials in the central gray matter show much smaller amplitudes compared with somatosensory cortical evoked potentials (SSEP). The strongest component is a negative wave after 70--100 msec. Longer conditioning stimulation of the lemniscal system inhibits late components in the median nerve evoked cortical potentials. On the contrary, stimulation of the nonspecific periaqueductal gray matter produces inhibition of early components of cortical SSEP together with facilitation of late components.     

Auditory evoked potentials to abrupt pitch and timbre change of complex tones: electrophysiological evidence of `streaming'?

Examination of the cortical auditory evoked potentials to complex tones changing in pitch and timbre suggests a useful new method for investigating higher auditory processes, in particular those concerned with `streaming' and auditory object formation. The main conclusions were: (i) the N1 evoked by a sudden change in pitch or timbre was more posteriorly distributed than the N1 at the onset of the tone, indicating at least partial segregation of the neuronal populations responsive to sound onset and spectral change; (ii) the T-complex was consistently larger over the right hemisphere, consistent with clinical and PET evidence for particular involvement of the right temporal lobe in the processing of timbral and musical material; (iii) responses to timbral change were relatively unaffected by increasing the rate of interspersed changes in pitch, suggesting a mechanism for detecting the onset of a new voice in a constantly modulated sound stream; (iv) responses to onset, offset and pitch change of complex tones were relatively unaffected by interfering tones when the latter were of a different timbre, suggesting these responses must be generated subsequent to auditory stream segregation.     

Functional organization of auditory projections to somatosensory cortical areas in cats

The functional properties of fibers transmitting auditory impulses to somatosensory areas S_I and S_II were studied in anesthetized and waking animals by the evoked potentials method. The thresholds of evoked potentials in areas S_I and S_II are 15–35 dB higher than those of evoked potentials in the auditory projection areas. Tonotopical localization is absent in somatic areas. Experiments on anesthetized animals showed that the spread of impulses relating to acoustic stimuli of different frequencies into areas S_I and S_II is effected through area A_I and its connections with the above zones. Another pathway probably also participates in the conduction of impulses from clicks. Analysis of the time constants of the first positive potential suggested that the interneuronal organization of auditory projections to area A_I is less complex than that of projections to the somatosensory areas. Comparison of amplitudes of evoked potentials of different projection zones in area S_I showed that the projection of the head receives more auditory impulses than the projection zone of the forelimbs, confirming the morphological data published previously.     

Genesis and functional role of cortical evoked potentials

Experiments on cats with recording electrodes implanted into the cranial bone showed that the evoked potential (EP) in the auditory cortex of the intact waking cat in response to clicks consists of five components with a total duration of up to 300 msec. Neurons of two types participate in the response to clicks: those with and without background activity. The former respond to clicks by various changes in background activity, the latter by one or several action potentials. The latent period of this response varies in different neurons from 6 to 250 msec. In response to clicks, several groups of neurons participate successively in the response, accounting for its long duration. From the beginning of the response, neurons of all cortical layers take part in it. At any moment of EP development, some neurons are in a state of excitation, others in a state of inhibition. About 80% of neurons responding to clicks respond before or during the initial electropositivity, 12% during the initial electronegativity, and only 8% during the late components of the EP. The importance of these findings is discussed relative to the question of the nature of the EP and of processes taking place in the brain after the arrival of an afferent volley.A. A Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neiofiziologiya, Vol. 2, No. 4, pp. 349–359, July–August, 1970.     

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.     

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.     

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.     

Age changes in latent periods of the human slow auditory evoked potential

With age regular changes take place in the latent periods of spikes of the slow auditory evoked potential. In particular, the latencies of the comparatively early waves (P₁, N₁, and P₂) become progressively shortened. Between 3–7 and 8–13 years the decrease is 50–60 msec, and later it is 25–35 msec. The latencies of the latest waves, especially P₃, N₃, and P₄, increase from 3–7 to 8–13 years by 35–65 msec. Later the latent period of the P₃ spike remains unchanged but the N₃ and P₄ waves disappear completely. Of all the components of the slow auditory evoked potential the most stable is the N₂ wave, the latent period of which decreases only very slightly with age. In children aged 3–7 years two wave complexes (P₁N₁P₂ and P₂N₂P₃) overlap frequently to form a single undifferentiated wave. This splits up into its components by 8 years of age. Long age changes in the shape and parameters of the slow auditory evoked potential are examined from the standpoint of the predominantly extralemniscal origin of this potential. On the basis of correlation discovered between the late waves of the evoked potential and the level of EEG synchronization it is postulated that the late waves of the slow evoked potential are formed with the participation of the nonspecific synchronizing system.Tbilisi State Postgraduate Medical Institute. Translated from Neirofiziologiya, Vol. 9, No. 1, pp. 3–10, January–February, 1977.     

Synaptic nature of the principal components of the evoked potential in the general cortex of the turtle forebrain

The nature of the principal components of the evoked potential of the general cortex of the turtle forebrain was studied in response to electrical stimulation of the contralateral optic nerve. Comparison of these components with postsynaptic potentials of the neurons of this structure showed that the four fast negative waves of the evoked potential correspond to fast EPSPs, which are independent of one another. The positive wave of the evoked potential is the sum of several IPSPs. The slow negative and, to some extent, the positive wave are a reflection of the slow EPSP. It is shown that early EPSPs are generated on portions of the apical dendrites which are further from the soma than those generating late fast EPSPs and also the IPSP and slow EPSP. Axo-somatic contacts are perhaps also concerned in the generation of the last-named potential.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 5, No.3, pp.261–271, May–June, 1973.     

The Enhancement of the N1 Wave Elicited by Sensory Stimuli Presented at Very Short Inter-Stimulus Intervals Is a General Feature across Sensory Systems

Background

A paradoxical enhancement of the magnitude of the N1 wave of the auditory event-related potential (ERP) has been described when auditory stimuli are presented at very short (<400 ms) inter-stimulus intervals (ISI). Here, we examined whether this enhancement is specific for the auditory system, or whether it also affects ERPs elicited by stimuli belonging to other sensory modalities.

Methodology and Principal Findings

We recorded ERPs elicited by auditory and somatosensory stimuli in 13 healthy subjects. For each sensory modality, 4800 stimuli were presented. Auditory stimuli consisted in brief tones presented binaurally, and somatosensory stimuli consisted in constant-current electrical pulses applied to the right median nerve. Stimuli were delivered continuously, and the ISI was varied randomly between 100 and 1000 ms. We found that the ISI had a similar effect on both auditory and somatosensory ERPs. In both sensory modalities, ISI had an opposite effect on the magnitude of the N1 and P2 waves: the magnitude of the auditory and the somatosensory N1 was significantly increased at ISI≤200 ms, while the magnitude of the auditory and the somatosensory P2 was significantly decreased at ISI≤200 ms.

Conclusion and Significance

The observation that both the auditory and the somatosensory N1 are enhanced at short ISIs indicates that this phenomenon reflects a physiological property that is common across sensory systems, rather than, as previously suggested, unique for the auditory system. Two of the hypotheses most frequently put forward to explain this observation, namely (i) the decreased contribution of inhibitory postsynaptic potentials to the recorded scalp ERPs and (ii) the decreased contribution of ‘latent inhibition’, are discussed. Because neither of these two hypotheses can satisfactorily account for the concomitant reduction of the auditory and the somatosensory P2, we propose a third, novel hypothesis, consisting in the modulation of a single neural component contributing to both the N1 and the P2 waves.     

Area 39 and 41 neurons sensitive to acoustic stimulation in the rat cortex: Polysensory properties

A comparative analysis of the polysensory properties of 102 neurons in areas 39 and 41 (the associative and auditory cortices, respectively) was performed in acute experiments on rats under chloralose-nembutal anesthesia. In the auditory cortex, the evoked potentials (EP) recorded from the surface of the above area in response to acoustic tonal, electrical cutaneous, and light stimulations almost always were distinguished by their shorter (4–5 msec) latency and higher amplitude. We studied neurons in both areas; their responses to the pure tones of various frequencies and to the stimulations of other modalities were compared. Bi- and polysensory neurons constituted 56.4% in area 39, and only 23% in area 41. The depth distribution of the responding neurons in areas 39 and 41 was different. Neurons with selective sensitivity to different frequencies of tonal signals were found in both areas. Usually monomodal neurons demonstrated selective properties in the auditory cortex, and 70% of them had a characteristic frequency. Over one-half of polymodal cells were frequency-selective in the associative cortex.Neirofiziologiya/Neurophysiology, Vol. 26, No. 3, pp. 223–229, May–June, 1994.