Statistical analysis of the parameters defining the cortical visual evoked potentials in the phases of the sleep-wake cycle

Visual evoked potential parameters (latencies, intervals of latencies and amplitudes) obtained by photic stimulation using a light-emitting diode implanted in the frontal sinus of cats were studied by statistical methods (analysis of variance) during the stages of wakefulness, slow sleep and paradoxical sleep. The results show: a) greater intraindividual homogeneity in all cases with special emphasis on the latencies; b) the greatest homogeneity of responses was found during slow sleep and paradoxical sleep stages; c) in relation to the influences exerted by the sleep-wakefulness cycle on the visual evoked potentials, the parameters most affected were those closely related to the secondary complex. We conclude, that latency, due to its great homogeneity, is the most useful parameter in this kind of experiments and secondly, that it is the secondary complex of the visual evoked potentials that is affected by the endogenous conditions of the subject (in our case the sleep-wakefulness cycle stage).     

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.     

Double evoked responses to the electrical stimulation of Shaffer's collaterals in CA1 hippocampal field

Double evoked responses to single current pulses applied to Shaffer's collaterals were observed in CAI hippocampal field in freely moving rats. The second response was an irregular population EPSP with constant latency sometimes accompanied by a population spike. The effect was observed in 22 of 54 tested rats (40.74%). In 10 of these 22 animals (45.45%) the second response was evoked by a weak testing stimulus of at least of a single collateral, in the remaining 12 rats the second response appeared to stimulation with increased current or after potentiation of the stimulated pathways. The second responses were very sensitive to the functional state of an animal and were recorded at the state of rest or during sleep. The latencies of the second response measured from the beginning of the first response were very close in different animals and with low intraindividual variability indicating that the same circuits are involved in its generation. It is suggested that hippocampus can support dynamic processes such as reverberation of signals.     

Recognition of joy, anger, and fear by face expression in humans

Behavioral and neurophysiological characteristics of a visual recognition of emotions of joy, anger, and fear were studied in 9 young healthy men and 10 women. It was shown that these emotions were identified by subjects with different rate and accuracy; significant gender differences in recognition of anger and fear were found. Recording of visual evoked potentials (VEP) from the occipital (O1/2), medial temporal (T3/4), inferior temporal (T5/6), and frontal (F3/4) areas revealed differences (related with the type of emotion) in the latencies of P150, N180, P250, and N350 waves and in the amplitude of VEP components with the latencies longer than 250 ms. These differences were maximally expressed in T3/4 derivation. The subjects could be divided in two groups. The first group was characterized by increased VEP latencies and higher amplitudes of VEP components later than 250 ms in response to anger (in comparison with other types of emotions). These phenomena were observed in all the derivations but were most pronounced in T3/4. In the second group, only late P250 and N350 components had shorter latencies during recognition of fear. VEP amplitude variations related with the type of emotions were insignificant and were recorded in the occipital and frontal areas. The two groups of subjects also differed in psychoemotional personality characteristics. It is suggested that primary recognition of facial expression takes place in the temporal cortical areas. A possible correlation of electrophysiological indices of emotion recognition with personality traits is discussed.     

Scalp-recorded oscillatory potentials evoked by transient pattern-reversal visual stimulation in man

Replicable oscillatory potentials, time-locked to pattern stimuli (9.0° central; counterphase reversal at 2.13 Hz) were dissociated from conventional, broad-band VEPs recorded in healthy volunteers at occipital scalp locations by high-pass digital filtering at 17.0–20.0 Hz. Nine consecutive wavelets were identified with a 56.4 ± 8.4 msec mean latency of the first replicable wavelet and mean peak-to-peak amplitude varying between 0.9 and 2.0 μV. The first 2 wavelets had significantly shorter latencies than wave N70 of unfiltered VEP, whereas the last 2 wavelets had longer latencies than N145. Latency and amplitude values varied as a function of contrast and spatial frequency of the stimulus, with shorter latencies and larger amplitudes at 60–90% contrast level and tuning of amplitude at 5.0 c/deg. All wavelets were correlated with wave P100 of unfiltered VEP, while a correlation with N70 of VEP was observed only for those wavelets with latencies in the range of wave P100. Two patients with documented brain lesions involving the visual system are described as examples of oscillatory responses occurring irrespective of filter bandpass and instead of the expected conventional VEP when the generation of these is interfered with by brain pathology. A substantial cortical contribution to the origin of the oscillatory response is conceivable. It is suggested that the oscillatory response to pattern-reversal stimulation reflects events in the visual system that are parallel to, and partly independent of, the conventional VEP, with potential application in research or for clinical purposes.     

The wake-sleep cycle in the cat following dorso-pontine lesions

Modifications in total amount, frequency and duration of episodes of wakefulness (W), drowsiness (D), slow sleep (SS) and paradoxal sleep (PS) were examined in cats with small unilateral lesion in the locus coeruleus (LC) and/or neighbouring structures. Throughout a 8 week period, 24 h weekly records were obtained from each cat. Control group with lesions in the dorsomedial pontine area showed a noticeable stability in sleep-wakefulness cycle (SWC). Significant decrease of W (with shortening of episodes) occurred during the whole postlesion period in the group with lesions in the lateral vestibular nucleus. The group with small lesions restricted to the dorsal-intermediate part of RPO and RPC nuclei had only acute SWC modifications consisting of a decrease of W and increase of PS. Finally, small lesions in LC complex that encroached to a limited extent neighbouring structures produced a maintained decrease of W (with shortening of episodes) and increase of PS (due to a major number of episodes). Results suggest a different role of dorsopontine region from other pontine areas in SWC mechanisms and give support to the hypothesis of a permissive role of LC in PS mechanisms.     

Conditioned modification of viscerosensory evoked potentials during sleep and wakefulness in cats

The effects of classic conditioning on the viscerosensory evoked potentials (EPs) were studied in twenty cats during wakefulness (W), slow-wave-sleep (SWS) and paradoxical sleep (PS). Four types of the experiment were performed on four groups of animals. Weak, non-painful stimulation of the small intestine or of the left splanchnic nerve was used as conditional stimulus (CS) in all experiments. A painful or non-painful shock on the left radial nerve served as unconditional stimulus (US) which followed the CS with a delay of 500 ms. In the first and second series of experiments, the CS was paired with non-painful or painful CS during W. In the third and fourth types of experiment, weak US was used and conditioning was done during SWS or PS. The evoked responses were recorded from the primary (SI) and secondary (SII) somatosensory and associative (AS) cortex, the thalamus (VPL), hypothalamus (HPT) and dorsal hippocampus (HPC). In each experiment, the stimulus pairings resulted in a complex electrographic conditional response (CR) which included an amplitude increase of the late components of EP's (early CR) and the development of a wave of 500 ms latency (delayed CR). In the second experiment, however, a behavioural CR (limb flexion) also appeared. All these CRs proved to be extinguishable. The recall of CR established during W was successful in SWS. The traces of CS-US pairings during SWS could, however, be elicited only in SWS. Both establishment and recall of CR were unsuccessful during PS. The possible mechanism of the effects originating from an interaction of conditioning and sleep on the viscerosensory inputs of the brain are discussed.     

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.     

The temporal frequency response function of pattern ERG and VEP: changes in optic neuritis

Steady-state pattern electroretinograms (PERGs) and visual evoked potentials (VEPs) in response to sinusoidal gratings (2 c/deg), sinusoidally counterphased at closely spaced temporal frequencies (TFs) between 4 and 28 Hz, were recorded from 11 patients with unilateral optic neuritis (ON; 11 affected eyes and 10 healthy fellow eyes) and 7 age-matched normal subjects (7 eyes). Amplitude and phase of responses' second harmonics were measured. Responses' apparent latencies were estimated from the rate at which phase lagged with TF. When compared to control values, mean PERG and VEP amplitudes of ON eyes were reduced (by about 0.4 log units) at both low (5–10 Hz) and high (16–20 Hz) TFs. Mean PERG amplitudes of fellow eyes were selectively reduced at low TFs (by about 0.3 log units). Mean PERG apparent latencies of both ON and fellow eyes were delayed (by 15 and 9 ms, respectively). Mean VEP apparent latency of ON eyes was delayed at both low and high TFs (by 24 and 30 ms, respectively), while that of fellow eyes was selectively delayed at high TFs (by 28 ms). The results in ON eyes indicate non-selective abnormalities of PERG and VEP generators responding at both low and high TFs. VEP TF losses may be in part accounted for by corresponding PERG losses. In the fellow eyes of patients, more selective PERG and VEP TF abnormalities may suggest differential impairment of retinal and postretinal subsystems responding better to low and high TFs (i.e. parvo-and magnocellular streams).     

侧脑室注射促甲状腺激素释放激素对大鼠睡眠—觉醒的影响

采用多导睡眠描记术研究了例脑室注射促甲状腺激素释放激素(TRH)对正常大鼠和去甲状腺大鼠睡眠-觉醒的影响。在正常大鼠,TRH引起觉醒增加,浅慢波睡眠(SWS_1)、深慢波睡眠(SWS_2)和总睡眠时间(TST)均减少,异相睡眠(PS)消失,SWS_1、SWS_2和PS的潜伏期均显著延长,给药后立即产生效应并在1h内达高峰。去甲状腺对大鼠的睡眠-觉醒无明显影响,注射TRH后引起的效应与正常大鼠相似。结果提示TRH有促进大鼠觉醒的作用,对各睡眠时相均有抑制作用,其作用部位可能在下丘脑以外的中枢结构。     

Effect of sleep stage on somatosensory evoked potentials by median nerve stimulation

The effects of sleep stage on early cortical somatosensory evoked potentials (SEPs) and short-latency components elicited by median nerve stimulation were studied in 12 normal volunteers. The latency of P13 in the awake stage was not significantly different from that in any sleep stage. The latencies of N16, N20 and P20 were significantly prolonged while the amplitude of N20 was decreased during the non-rapid eye movement (NREM) sleep stage. P22, P23 and N24 components showed double peaks (P23a, P23b, N24a, N24b) during the NREM sleep stage in 6 subjects, while N24 showed a single peak and only P22 and P23 showed double peaks in 5 other subjects. The latencies and morphologies of SEPs during rapid eye movement sleep stage were almost the same as those during the awake stage. These findings suggest that NREM sleep affects the latency, amplitude and morphology of N16 and early cortical components.     

Human growth hormone release is decreased during sleep in temporal isolation (free-running)

Growth hormone (hGH) secretion was measured during sleep in 10 healthy male subjects isolated from all time cues. HGH concentrations following sleep onset were compared between scheduled sleep episodes (entrainment) and self-selected sleep episodes (free-running). Peak sleep-related hGH values were decreased significantly during free-running. The duration of the first slow wave sleep (SWS) episode and the latency to the first REM sleep episode also decreased significantly during free-running. The latencies from sleep onset to SWS and to peak hGH secretion did not differ between entrainment and free-running. These results suggest that sleep-related hGH secretion begins 'on time' during free-running, but is terminated earlier. Thus, while sleep onset facilitates hGH release, the timing of other stages of sleep such as REM may alter the magnitude of sleep-related hGH secretion.     

Photic evoked responses in the brain stem of the rat]

The appearance of photic evoked responses in various structures of the reticular formation (RF) and the thalamus was investigated in freely moving rats. Photic evoked potentials (PhEP) were recorded in all the nuclei tested. The PhEP are different in shape, amplitude and latency; they can be classified as a "primary type" with small amplitudes and without latency differences from the PhEP of the visual cortex (VC) and as a "secondary type" with large amplitudes and latency differences from the PhEP of the VC. The "primary type" was observed in thalamic, pontine and bulbar structures but the "secondary type" in posterior-thalamic and mesencephalic structures. Photic afterdischarges (PhNE) and photic recruitment (PhR) were recorded in most of the nuclei tested. These PhNE and PhR have a correlation in their frequency and peak-latency to the PhNE and PhR of the VC. It is discussed that a great part of visual information is transferred to the brain stem through the Corpus geniculatum laterale (CGL) and the VC.     

Differential action upon sleep states of ventrolateral and central areas of pontine tegmental field

In order to study oral pontine mechanisms of the sleep- wakefulness cycle (SWC), modifications in the total amount, frequency, and duration of episodes of wakefulness (W), drowsiness (D), slow sleep (SS) and paradoxical sleep (PS) together with modifications in the hourly distribution of both sleep states were analyzed in 15 adult cats. Three animals were used as sham-operated controls. Six of them, Group I, received unilateral lesions in ventral and lateral areas of the nucleus reticularis pontis oralis (RPO) and rostral nucleus reticularis pontis caudalis (RPC). The remaining 6 cats, Group II, had unilateral lesions in the central part of the same nuclei. After ventrolateral lesions (Group I) decrease of PS and SS occurred, but only PS changes reached statistically significant values; while, on the contrary, a significant increase of SS and PS followed central lesions (Group II). Hourly distribution analysis indicated that in Group I decrease of both sleep states took place mainly at night, while in Group II increase of both SS and PS occurred during day. These results suggest a complex and non- uniform influence of the pontine tegmental area on SWC mechanisms. Effects obtained after unilateral lesions, precisely located in ventral and lateral or central parts, point to the existence of two functionally distinct, although almost overlapping, systems, at this level.     

Dynamics of potentials evoked in the auditory pathway and reticular formation of the cat. Studies during waking and sleep stages

Averaged evoked potentials in the inferior colliculus (IC), medial geniculate nucleus (MG) and reticular formation (RF) of chronically implanted and freely moving cats were measured using auditory step functions in the form of tone bursts of 2000 Hz. The most prominent components of the AEP of the inferior colliculus were a positive wave of 13 msec and a negative wave of 40–55 msec latency. The AEP of the medial geniculate nucleus was characterized by a large negative wave peaking at 35–40 msec. During spindle sleep and slow wave sleep stages changes in the AEPs of both nuclei occured.Transient evoked responses of the inferior colliculus, medial geniculate nucleus and reticular formation were transformed to the frequency domain using the Laplace transform (one sided Fourier transform) in order to obtain frequency characteristics of the systems under study. The amplitude characteristics of IC, MG. and RF obtained in this way revealed maxima in alpha (8–13 Hz), beta (18–35 Hz) and higher frequency (50–80 Hz) ranges. During spindle sleep stage a maximum in the theta frequency range (3–8 Hz) and during slow wave sleep maximum in the delta (1–3 Hz) frequency range appeared in the amplitude characteristics of these nuclei.The amplitude characteristics of the inferior colliculus and medial geniculate nucleus were compared with the amplitude characteristics of other brain structures. The comparison of AEPs and amplitude frequency characteristics obtained using these AEPs reveals that the existence of a number of peaks (waves) with different latencies in the time course does not necessarily indicate the existence of different functional structures or neural groups giving rise to these waves. The entire time course of evoked potentials and not the number and latencies of the waves, carries, the whole information concerning different activities and frequency selectivities of brain structures.Supported by Turkish Scientific and Technical Research Council Grant TAG-266.Presented in Part at the VIIIth International Congress of Electroencephalography and Clinical Neurophysiology in Marseilles, September 1–7, 1973.     

Pre-stimulus spectral EEG patterns and the visual evoked response

The relationship between the latencies and amplitudes of the N1 and P2 components of the visual evoked potential (VEP) and the psychophysiological state of the brain immediately preceding the time of the stimulus has been investigated in 7 male subjects. Power spectral measures in the delta, theta, alpha and beta bands of the 1 sec pre-stimulus EEG were used to assess the brain state, and low intensity flashes, delivered randomly between 2 and 6 whole seconds, were used as the stimuli. Trials were ranked separately according to the relative amounts of pre-stimulus power in each EEG band and were partitioned into groups by an equal pre-stimulus spectral power criterion. Averaged EPs were computed from these groups and multiple regression analysis was used to relate pre-stimulus spectral power values to EP features. Five of the 7 subjects displayed consistent increases in N1-P2 amplitude as a function of increasing pre-stimulus relative alpha power. The between-subjects effect of pre-stimulus EEG on N1 latency was small, but was moderate for P2 latency (both significant). Both N1 and P2 latency were found to decrease with increasing amounts of pre-stimulus relative delta and theta power.     

Neurophysiological effects of recombinant interferon-gamma and -alpha in man

Treatment of malignant tumors with interferon (IFN) is in some patients accompanied by serious neurological side effects. The present study assessed neurophysiological changes in spontaneous EEG activity, visual-evoked cortical potentials (VEPs), and brainstem auditory-evoked potentials (BAEPs) during IFN-gamma or IFN-alpha therapy in 9 patients. In addition, blood pressure, heart rate and body temperature were monitored. In all sessions under IFN, the latency of the P100 component of the VEP was shortened as compared to baseline conditions. IFN also reduced latencies of BAEP components, and diminished amplitudes of the spontaneous EEG activity within the alpha and beta frequency band. These latter effects were somewhat less consistent than those on VEPs. The major neurophysiological changes appeared to be similar for IFN-gamma and IFN-alpha. The results are in accord with an excitatory effect of IFN on central nervous activity. The magnitude of changes excludes a neurotoxicity of IFN-gamma or IFN-alpha at the doses used in this study.     

Facilitation of the diaphragm response to transcranial magnetic stimulation by increases in human respiratory drive.

The human respiratory neural drive has an automatic component (bulbospinal pathway) and a volitional component (corticospinal pathway). The aim of this study was to assess the effects of a hypercapnia-induced increase in the automatic respiratory drive on the function of the diaphragmatic corticospinal pathway as independently as possible of any other influence. Thirteen healthy volunteers breathed room air and then 5 and 7% hyperoxic CO2. Cervical (cms) and transcranial (tms) magnetic stimulations were performed during early inspiration and expiration. Transdiaphragmatic pressure (Pdi) and surface electromyogram of the diaphragm (DiEMG) and of the abductor pollicis brevis (apbEMG) were recorded in response to cms and tms. During inspiration, Pdi,cms was unaffected by CO2, but Pdi,tms increased significantly with 7% CO2. During expiration, Pdi,cms was significantly reduced by CO2, whereas Pdi,tms was preserved. DiEMG,tms latencies decreased significantly during early inspiration and expiration (air vs. 5% CO2 and air vs. 7% CO2). DiEMG,tms amplitude increased significantly in response to early expiration-tms (air vs. 5% CO2 and air vs. 7% CO2) but not in response to early inspiration-tms. DiEMG,cms latencies and amplitudes were not affected by CO2 whereas 7% CO2 significantly increased the apbEMG,cms latency. The apbEMG,tms vs. apbEMG,cms latency difference was unaffected by CO2. In conclusion, increasing the automatic drive to breathe facilitates the response of the diaphragm to tms, during both inspiration and expiration. This could allow the corticospinal drive to breathe to keep the capacity to modulate respiration in conditions under which the automatic respiratory control is stimulated.     

γ-氨基丁酸、荷包牡丹碱和L-谷氨酸钠对猫和家兔同侧和对侧图形视觉诱发电位的影响

在猫和家兔大脑半球一侧视区17/18交界处施加γ—氨基丁酸(GABA)、荷包牡丹碱和L—谷氨酸钠,以及用氯化钾和冷冻阻遏的方法,记录对侧和同侧皮层相应处图形视觉诱发电位(PVEP)的变化。讨论了GABA、荷包牡丹碱和L—谷氨酸钠对猫和兔的对侧和同侧PVEP的影响。     

Dynamics of neuronal activity in the cingulate gyrus during the sleep-wake cycle

Dynamics of neuronal activity in the cingulate gyrus (CG) was investigated during the sleep-wake cycle (SWC) of free-ranging cats. The highest activity rate was found to occur in 65.4% of neurons during active emotional consciousness (EC) and the so-called emotional phase of paradoxical sleep (PS), while firing rate decreased during passive consciousness (PC) and slow-wave sleep (SS). Peak firing rate was observed during SS in 15% of neurons, while divisions of activity between stages of the SWC remained the same in 19.6%. In addition, discharge patterns in 75.2% of neurons changed consistently in step with phase shifts. More particularly, neurons fired during EC and PS with single action potentials with a more or less even time distribution, whereas a burst-phase activity pattern occurred in the course of SS. Firing rate declined (in 42.6% of units) or remained unchanged (in 50.4%) in the majority of CG neurons (even amongst those manifesting highest activity during EC and PS) as episodes of isolated EEG arousal developed (whether in the context of SS or as PS wore off). This would indicate that the CG actually contributes to shaping the behavioral state of consciousness. The CG, therefore, as one of the higher divisions of the limbic system, must play a major part in controlling the basic mechanisms of the SWC, as well as emotionally motivated processes evolving in the cycle.I. S. Beritashvili Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 21, No. 6, pp. 832–840, November–December, 1989.