Phasic brain activity related to the onset of rapid eye movements during rapid eye movement sleep: study of event-related potentials and standardized low-resolution brain electromagnetic tomography.

The function of rapid eye movements (REMs) during REM sleep is still a matter that is open to debate. In a previous study, we found positive brain potential (P200r) time-locked to the onset of REMs. This potential was not observed during saccades of wakefulness. In this study, we estimated the electrical generation of this potential to investigate the phasic brain activity related to REMs. Data were collected in a sleep laboratory from nine healthy university students. REMs during REM sleep were recorded during natural nocturnal sleep. Event-related potential time-locked to the onset of REMs were averaged. Standardized low-resolution brain electromagnetic tomography (sLORETA) was used to identify the current sources of P200r. The results showed that P200r have neuronal generators in the left premotor area, left primary motor and sensory cortices, left inferior parietal lobule and bilateral occipital areas (precuneus, cuneus and lingual gyrus). All these areas are known to contribute to visuomotor processing. These phasic brain activities might play a key role in explaining the function of REMs during REM sleep.

Activation of fast sleep spindles at the premotor cortex and parietal areas contributes to motor learning: a study using sLORETA.

OBJECTIVE: The present study examined whether slow and/or fast sleep spindles are related to visuomotor learning, by examining the densities of current sleep spindle activities. METHODS: Participants completed a visuomotor task before and after sleep on the learning night. This task was not performed on the non-learning night. Standard polysomnographic recordings were made. After the amplitudes of slow and fast spindles were calculated, sLORETA was used to localize the source of slow and fast spindles and to investigate the relationship between spindle activity and motor learning. RESULTS: Fast spindle amplitude was significantly larger on the learning than on the non-learning nights, particularly at the left frontal area. sLORETA revealed that fast spindle activities in the left frontal and left parietal areas were enhanced when a new visuomotor skill was learned. There were no significant learning-dependent changes in slow spindle activity. CONCLUSIONS: Fast spindle activity increases in cortical areas that are involved in learning a new visuomotor skill. The thalamocortical network that underlies the generation of fast spindles may contribute to the synaptic plasticity that occurs during sleep. SIGNIFICANCE: Activity of fast sleep spindles is a possible biomarker of memory deficits.

Temporal coupling of rapid eye movements and cerebral activities during REM sleep.

OBJECTIVES: We investigated event-related potentials time locked to the onset and offset of rapid eye movements during rapid eye movement (REM) sleep. METHOD: Nine healthy university students participated in this study. Data were collected in a sleep laboratory. Rapid eye movements during REM sleep were recorded during natural nocturnal sleep. Saccades during wakefulness were recorded during a visually triggered task. Event-related potentials were averaged, time-locked to the onset and offset of eye movements. RESULTS: During REM sleep, a lambda-like response occurred over the occipital region, time-locked to the offset of rapid eye movements (similar to what occurs during wakefulness). Moreover, we found that a positive potential (P200r) occurred at about 200ms, with the maximal amplitude over the central region and time-locked to the onset of rapid eye movements during REM sleep; this potential was not observed during wakefulness. CONCLUSIONS: During REM sleep, the P200r occurs with the start of rapid eye movements, and then the lambda-like response occurs after termination of the movements. SIGNIFICANCE: We demonstrated temporal coupling of rapid eye movements and cerebral activities during REM sleep. These activities might provide a useful basis for future investigations of brain functions during REM sleep.

Neural generators of brain potentials before rapid eye movements during human REM sleep: a study using sLORETA.

OBJECTIVE: Brain activity preceding rapid eye movements (REM) during human REM sleep has remained poorly understood. Slow negative brain potential (pre-REM negativity) appears before REMs. Current sources of this potential were investigated to identify brain activity immediately preceding REMs. METHODS: In this study, 22 young healthy volunteers (20-25 years old) participated. Polysomnograms were recorded during normal nocturnal sleep. Brain potentials between 200ms before and 50ms after the onset of REMs and pseudo-triggers (3000ms before the onset of REMs) were averaged. Standardized low-resolution brain electromagnetic tomography (sLORETA) was used to estimate current sources of pre-REM negativity. RESULTS: Pre-REM negativity appeared with the maximal amplitude at right prefrontal sites immediately before REMs. However, this negativity did not appear before pseudo-triggers. Current sources of the pre-REM negativity were estimated in the ventromedial prefrontal cortex, uncus, insula, anterior cingulated cortex, basal forebrain, parahippocampal gyrus, premotor cortex and frontal eye field. CONCLUSIONS: The pre-REM negativity reflects brain activity coupled with the occurrence of REMs. Results of this study suggest that emotion, memory, and motor-related brain activity might occur before REMs. SIGNIFICANCE: Pre-REM negativity is expected to be a psychophysiological index for elucidating functions of REM sleep.

Fast sleep spindle (13-15 hz) activity correlates with sleep-dependent improvement in visuomotor performance.

STUDY OBJECTIVES: The relationship between memory enhancement and fast (13-16 Hz) versus slow (10-13 Hz) spindle activity during sleep was investigated. DESIGN: Standard polysomnographic recordings were conducted during an adaptation, control nonlearning, and learning night. Automatic spindle detection and measurement was utilized with visual confirmation. SETTING: Participants slept in individual, temperature-controlled bedrooms in a sleep laboratory. PARTICIPANTS: Twelve healthy student volunteers (9 women and 3 men, mean age: 22.3 years) participated. INTERVENTIONS: On the learning night, participants completed a presleep learning session on a modified version of mirror-tracing task followed by a postsleep test session. No learning or test sessions were performed on the adaptation and nonlearning nights. MEASUREMENTS AND RESULTS: Tracing time was reduced by 6.4 seconds (20.6% +/- 2.07%) from the presleep to the postsleep session. Mean amplitude and duration of fast spindles was greater on the learning night than on the nonlearning night (both P values < 0.05). Skill improvement and fast-spindle activity were positively correlated (density [r = 0.76, P < 0.01], amplitude [r = 0.69, P < 0.05], and duration [r = 0.67, P <0.05]). Significant correlations between fast-spindle activity and mirror-tracing performance were also evident for the nonlearning night. There was no significant relationship between mirror-tracing performance and slow-spindle activity on any night. CONCLUSIONS: The thalamocortical network underlying fast-spindle generation may contribute to or reflect plasticity during sleep.

Visual emotional context modulates brain potentials elicited by unattended tones.

To examine whether brain electrical responses to environmental stimuli were influenced by emotional contexts, event-related potentials (ERPs) elicited by nonstartle probe tones were recorded from 13 student volunteers while they were viewing emotionally positive, neutral, and negative slides of the International Affective Picture System. The auditory stimuli consisted of high-deviant (2000 Hz, p=.08), low-deviant (1050 Hz, p=.08), and standard (1000 Hz, p=.84) tones with a mean onset-to-onset interval of 600 ms. Participants were told to ignore the tones. High-deviant tones elicited a larger N1 (peaking around 100 ms) when participants were viewing negative slides than when viewing positive slides. The amplitude of the P2 elicited by standard tones (peaking around 170 ms) was smaller when participants were viewing positive slides than when viewing negative and neutral slides. The amplitude of the mismatch negativity (150-200 ms) tended to reduce during positive slide presentation, but this difference appeared to be due to reduction of the P2 elicited by standard tones. These findings suggest that visually induced emotional states have a sequential effect on auditory information processing, in that the influence of negative emotion appears at an earlier stage than that of positive emotion.

Anterior N2 predicts subsequent viewing time and interest rating for novel drawings.

Event-related brain potentials (ERPs) in response to novel pictures and subsequent visual exploration were examined. Eighteen undergraduates viewed 120 novel drawings as long as they wished. ERPs were calculated separately for drawings that were viewed longer or shorter than the median viewing time of each participant. The drawings viewed longer elicited a larger anterior negativity (N2) peaking at 245 ms than the shorter viewed drawings. This effect disappeared and the overall amplitude of the N2 decreased when the same set of drawings was presented again in a subsequent interest-rating session. Drawings rated as more interesting elicited a larger N2 than did drawings rated as less interesting. This study demonstrates a type of anterior N2 that is sensitive to stimulus unfamiliarity or difficulty in encoding, which reflects a conflict between stimulus input and existing knowledge and prompts further recognition processes and visual exploration.

Effect of voluntary attention on auditory processing during REM sleep.

STUDY OBJECTIVES: The study investigates whether there is an effect of voluntary attention to external auditory stimuli during rapid eye movement (REM) sleep in humans by measuring event-related potentials (ERPs). DESIGN: Using a 2-tone auditory-discrimination task, a standard 1000-Hz tone and a deviant 2000-Hz tone were presented to participants when awake and during sleep. In the ATTENTIVE condition, participants were requested to detect the deviant stimuli during their sleep whenever possible. In the PASSIVE sleep condition, participants were only exposed to the tones. ERPs were measured during REM sleep and compared between the 2 conditions. SETTING: All experiments were conducted at the sleep laboratory of Hiroshima University. PARTICIPANTS: Twenty healthy university student volunteers. INTERVENTIONS: N/A. MEASUREMENTS AND RESULTS: In the tonic period of REM sleep (the period without REM), P200 and P400 were elicited by deviant stimuli, with scalp distributions maximal at central and occipital sites, respectively. The P400 in REM sleep showed larger amplitudes in the ATTENTIVE condition, whereas the P200 amplitude did not differ between the 2 conditions. No effects on ERPs due to attention were observed during stage 2 sleep. CONCLUSIONS: The instruction to pay attention to external stimuli during REM sleep influenced the late positive potentials. Thus electrophysiologic evidence of voluntary attention during REM sleep has been demonstrated.

Self-awakening, sleep inertia, and P3 amplitude in elderly people.

It is well known that sleepiness is sometimes experienced in the afternoon. A short afternoon nap is thought to be effective in reducing sleepiness. However, sleep inertia occurs even after a short nap, and this could be a major risk factor for injuries from falling by the elderly. In the present study, the effect of self-awakening on sleep inertia after a 20-min. nap was examined. Nine participants (M=74.1 yr., SD=5.0 yr.) took part in the three experimental conditions: the self-awakened nap, the forced-awakened nap, and the control (no-nap) conditon. Analysis showed sleepiness and performance after the nap significantly improved compared with the control condition. P3 amplitude tended to be larger after self-awakening than after forced-awakening. The present study indicates a 20-min. nap reduces afternoon sleepiness, and the application of self-awakening may contribute to higher arousal after a nap taken by this elderly group.

Brain potentials before and after rapid eye movements: an electrophysiological approach to dreaming in REM sleep.

STUDY OBJECTIVE: This study examined hypotheses regarding dreaming in rapid eye movement (REM) sleep by comparing brain potentials related to rapid eye movements in REM sleep with those in wakefulness. DESIGN: Within participants. SETTING: Data were collected in a sleep laboratory. PARTICIPANTS: Fifteen healthy university students who reported having dreams frequently. MEASUREMENTS AND RESULTS: Rapid eye movements in REM sleep were recorded during natural nocturnal sleep. Saccades in wakefulness were recorded during a self-paced visual search task. The presaccadic negativity before and the lambda response after eye movements were examined. It was assumed that the presaccadic negativity reflects voluntary readiness activity before eye movements, and the lambda response reflects visual information processing after saccades in wakefulness. Brain potentials were averaged, time-locked to the onset and offset of eye movements for the presaccadic negativity and the lambda response, respectively. In wakefulness, the presaccadic negativity occurred at the centroparietal site. However, no presaccadic negativity was found during REM sleep. Lambda-like responses (P1r, P2r) were observed in REM sleep over the parietooccipital site, as were the lambda responses (P1, P2) in wakefulness. CONCLUSIONS: The finding that rapid eye movements are initiated without preparation but elicit some neural activity in the cortical visual area suggests that rapid eye movements may trigger dream images.

Examination of the first-night effect during the sleep-onset period.

STUDY OBJECTIVES: The present study examined the first-night effect during the sleep-onset period using the 9 electroencephalogram stage scoring system. DESIGN: After a week of monitoring sleep-wake habits with sleep diaries and wrist actigraphy, polysomnography recording was made for 3 consecutive nights. SETTING: Participants slept in their own private, individual, temperature-controlled bedroom in a sleep laboratory at the university. PARTICIPANTS: Eleven healthy student volunteers (5 women and 6 men, 21 to 25 years old, mean 22.7 years) who had no experience sleeping in a laboratory participated in the study. INTERVENTIONS: N/A. MEASUREMENTS AND RESULTS: The electroencephalogram during the sleep-onset period was scored manually for every 5-second epoch into 9 electroencephalogram stages. Latencies of the electroencephalogram stages were delayed on the first night, especially during the alpha-wave intermittent stages. The average time of the alpha-wave train, intermittent (> 50%) and the electroencephalogram flattening stage increased on Night 1. Stage changes among these stages also increased on Night 1. In contrast, stage changes between the alpha-wave intermittent stage (< 50%) and the theta-wave stage increased on Night 3. CONCLUSIONS: Alpha-wave activity increased on Night 1, demonstrating that the activity of the wake-promoting system during the sleep-onset period was enhanced on the first night. From the second to the third night, the alpha-wave intermittent stage jumped to the theta-wave stage, omitting electroencephalogram flattening, suggesting that the electroencephalogram flattening stage is unlikely to appear during stable sleep-onset period. This is the first study to demonstrate the detail of the first-night effect during the sleep-onset period.

Effects of hypnagogic imagery on the event-related potential to external tone stimuli.

STUDY OBJECTIVES: The purpose of this study was to examine the influence of hypnagogic imagery on the information processes of external tone stimuli during the sleep onset period with the use of event-related potentials. DESIGN: Event-related potentials to tone stimuli were compared between conditions with and without the experience of hypnagogic imagery. To control the arousal level when the tone was presented, a certain criterion named the electroencephalogram stage was used. Stimuli were presented at electroencephalogram stage 4, which was characterized by the appearance of a vertex sharp wave. SETTING: Data were collected in the sleep laboratory at Hiroshima University. PARTICIPANTS: Eleven healthy university and graduate school students participated in the study. INTERVENTIONS: N/A. MEASUREMENTS AND RESULTS: Experiments were performed at night. Reaction times to tone stimuli were measured, and only trials with shorter reaction times than 5000 milliseconds were analyzed. Electroencephalograms were recorded from Fz, Cz, Pz, Oz, T5 and T6. There were no differences in reaction times and electroencephalogram spectra between the conditions of with and without hypnagogic imagery. These results indicated that the arousal levels were not different between the 2 conditions. On the other hand, the N550 amplitude of the event-related potentials in the imagery condition was lower than in the no-imagery condition. CONCLUSIONS: The decrease in the N550 amplitude in the imagery condition showed that experiences of hypnagogic imagery exert some influence on the information processes of external tone stimuli. It is possible that the processing of hypnagogic imagery interferes with the processing of external stimuli, lowering the sensitivity to external stimuli.

Recuperative power of a short daytime nap with or without stage 2 sleep.

STUDY OBJECTIVES: The recuperative effect of a nap of less than 30 minutes has been confirmed. Such naps consist mainly of stages 1 and 2 sleep. The present study examined whether sleep stage 1 or 2 contributed to the recuperative effect of a short nap. DESIGN: Repeated-measurement within-subject design. After sleep was restricted to 1.5 hours less than their usual nocturnal sleep, participants took a rest (No-nap condition) or a nap at 2:00 PM. In the nap condition, they were awakened after 5 minutes of stage 1 sleep (S1-nap condition) or 3 minutes after stage 2 sleep appeared (S2-nap condition). SETTING: University sleep laboratory. PARTICIPANTS: Ten healthy university students (aged 19 to 24 years). MEASUREMENTS: Subjective mood, performance on visual detection and symbol-digit substitution tasks, and the number of slow eye movements during a performance task were measured before and after the nap or rest. RESULTS: In the No-nap condition, subjective mood and performance deteriorated, and Slow eye movements increased during mid-afternoon, suggesting that the post-lunch dip occurred. In contrast, subjective alertness and performance improved and slow eye movements rarely occurred in the S2-nap condition. Although subjective sleepiness and fatigue improved, performance deteriorated and slow eye movements increased in the S1-nap condition. CONCLUSION: A daytime short nap containing 3 minutes of stage 2 sleep has recuperative effects, whereas these effects are limited following only stage 1 sleep.

Self-awakening prevents acute rise in blood pressure and heart rate at the time of awakening in elderly people.

Self-awakening, waking up at a designated time decided before sleeping, could prevent failure in the blood circulation vessel system such as heart attack, acute increases in heart rate or blood pressure upon waking. Previous research showed that anticipatory changes occurred in heart rate prior to awakening from a short nap by means of self-awakening for young participants. However, the effects of self-awakening remained unclear for elderly people. The present study examined the effects of self-awakening on heart rate and blood pressure in a short afternoon nap (20 min) among the elderly. Nine participants [74.1 (SD = 5.01) years old] underwent both self-awakening and forced-awakening conditions. In the self-awakening condition, it was revealed that blood pressure gradually increased before the scheduled time of awakening, and that heart rate did not show a rapid increase at arousal. In contrast, forced-awakening induced acute increases in both heart rate and blood pressure. These results suggest that self-awakening facilitates a more smooth transition from sleep to wakefulness via autonomic activation before the time of self-awakening.

Level of interest in video clips modulates event-related potentials to auditory probes.

This study examined whether a participant's level of interest in visual materials could be assessed by event-related potentials to auditory probe stimuli. Twelve young adults performed an auditory target detection task while viewing either interesting or less interesting (neutral) silent video clips. The auditory probe stimuli consisted of target (2000 Hz, p=0.15), standard (1800 Hz, p=0.70), and nontarget deviant (500 Hz, p=0.15) tones. Button press responses to target tones were required. Both target and deviant tones elicited a large P3 wave, the amplitude of which was smaller while participants were viewing interesting video clips than neutral video clips or simple still images (control condition). The amplitude reduction of the P3 to deviant tones was more prominent than that of the P3 to target tones. The difference between the neutral and control conditions was significant only for the deviant P3. The three-tone probe task using perceptually deviant, nontarget stimuli may be a useful objective method to assess how strongly a visual material attracts the viewer's attention.

Lack of presaccadic positivity before rapid eye movements in human REM sleep.

Differences between oculomotor control of rapid eye movements (REMs) in REM sleep and that of saccades in wakefulness were examined electrophysiologically in human adults. Fourteen healthy young volunteers participated in the study. Brain potentials were recorded from the scalp and time-locked to the onsets of saccades and REMs during a visually triggered saccade task and natural nocturnal sleep. In wakefulness, presaccadic positivity (PSP) appeared at centro-parietal sites starting about 150 ms before saccades. In REM sleep, no PSP was found but a slow negative potential (pre-REM negativity: PRN) appeared at the prefrontal sites. The findings suggest that the generation of REMs does not involve the cortical process reflected in the PSP but is associated with a different neural process reflected in the PRN.

Short nap versus short rest: recuperative effects during VDT work.

The effects of a 20-min nap during 2 h of visual display terminal (VDT) work were examined. Ten young healthy adults took a 20-min nap or a 20-min rest I h after VDT work, followed by another 1 h of VDT work. A 20-min rest temporarily restored subjective sleepiness, but it deteriorated during the additional 1 h of work. In contrast, a 20-min nap maintained subjective alertness and performance level at a higher level and mental fatigue at a lower level for the additional 1 h of work. These results suggest that a short nap would be useful to both fatigue recovery and fatigue prevention during continuous VDT work. The present findings may provide a new work/rest strategy.

Subjectively optimal delay in computer response to a user's voluntary action.

It is typically assumed that the shorter the delay in a computer response to a user's action is, the better. In a study examining this assumption, 16 university students were asked to judge the timing of occurrence of either an auditory or a visual stimulus following a voluntary mouse button-press with various durations of delay from 0 to 400 msec. Both a category scaling and an adjustment method of assessment were used to specify the optimal delay with which the user felt most comfortable. When the delay was 50 msec. or shorter, both auditory and visual responses were judged as too fast to be a consequence of the action. The responses were judged as too slow when the delay was 200 msec. or longer for auditory and 300 msec. or longer for visual stimuli. Analysis for the adjustment method indicated that the optimal delay was within 100 to 200 msec. and was longer for visual than for auditory responses. The present study suggests that immediate responses from a computer are not always comfortable and that the insertion of a moderate delay interval may increase the user's sense of control.

The alerting effects of caffeine, bright light and face washing after a short daytime nap.

OBJECTIVE: The present study examined whether the combination of a short daytime nap with caffeine, bright light and face washing was effective against mid-afternoon sleepiness. METHODS: Ten young healthy adults participated in 5 experimental conditions; those experiments were-Nap only: taking a 20 min nap; Caffeine+Nap: taking 200 mg of caffeine followed by a nap; Nap+Bright-light: being exposed to 2000 lx of bright light for 1 min immediately after napping; Nap+Face-washing: washing their faces immediately after napping; and No-Nap: taking a rest without sleep. These naps were taken at 12:40 hours. The subjects engaged in computer tasks for 15 min before napping and for 1 h after napping. RESULTS: Caffeine+Nap was the most effective for subjective sleepiness and performance level; its effects lasted throughout 1 h after napping. Nap+Bright-light was comparable with Caffeine+Nap, except for performance level. Nap+Face-washing showed mild and transient effects, however, it suppressed subjective sleepiness immediately after napping. CONCLUSIONS: The effects of a short nap against mid-afternoon sleepiness could be enhanced by combining caffeine intake, exposure to bright light, or face washing. SIGNIFICANCE: The present study would provide effective countermeasures against mid-afternoon sleepiness and sleepiness related accidents.

The effects of self-awakening on heart rate activity in a short afternoon nap.

OBJECTIVES: This study examined whether anticipatory changes exist in heart rate prior to awakening from a nap by means of self-awakening. The effects of self-awakening on sleepiness after the short nap were also studied. METHOD: Nine students participated in 3 experimental conditions: (1) the control condition, in which participants watched television instead of taking a short nap; (2) the self-awakening condition, in which participants tried to wake up approximately 15 min after 'lights off' (criterion range: 15+/-5 min) and (3) the forced-awakening condition, in which participants were instructed to sleep for 30 min, but were awoken by the experimenter after 15 min. RESULTS: In the self-awakening condition, heart rate gradually increased approximately 3 min before awakening. The error response ratio of the auditory-oddball task and the duration of doze time during the task were less after both types of nap conditions than in the control condition. Subjective sleepiness, which is measured after awakening, was lowest in the self-awakening condition. CONCLUSIONS: The results of this experiment suggest that self-awakening prepares autonomic activation that facilitates a more smooth transition from sleep to awakening, and reduces sleepiness after naps.