Hemispheric lateralization of the EEG during wakefulness and REM sleep in young healthy adults

EEG recordings confirm hemispheric lateralization of brain activity during cognitive tasks. The aim of the present study was to investigate spontaneous EEG lateralization under two conditions, waking and REM sleep. Bilateral monopolar EEG was recorded in eight participants using a 12-electrode montage, before the night (5 min eyes closed) and during REM sleep. Spectral analysis (0.75-19.75 Hz) revealed left prefrontal lateralization on total spectrum amplitude power and right occipital lateralization in Delta activity during waking. In contrast, during REM sleep, right frontal lateralization in Theta and Beta activities and right lateralization in occipital Delta activity was observed. These results suggest that spontaneous EEG activities generated during waking and REM sleep are supported in part by a common thalamo-cortical neural network (right occipital Delta dominance) while additional, possibly neuro-cognitive factors modulate waking left prefrontal dominance and REM sleep right frontal dominance.     

Frontal electroencephalogram alpha asymmetry during sleep: stability and its relation to affective style

Electroencephalogram (EEG) alpha (8-12 Hz) asymmetries were collected from the mid-frontal and central regions during presleep wakefulness and Stage 1, Stage 2, and rapid eye movement (REM) sleep in 11 healthy right-handed participants who were free of psychiatric, neurological, and sleep problems. The authors found significant correlations between presleep wakefulness and different stages of sleep in the frontal, but not central, EEG alpha asymmetry measure. The strongest correlation was between presleep waking and REM sleep, replicating and extending relation earlier work to a normal population. The high degree of association between presleep waking and REM sleep may be a result of high cortical activation common to these states and may reflect a predisposition to different styles of emotional reactivity.     

Variations in EEG coherence as an index of the affective content of dreams from REM sleep: relationships with face imagery

EEG coherence was examined in relation to four measures of socioemotional dream content, including a new measure--the proportional representation of a character's face. Twenty-four healthy subjects, recorded for sleep stages and EEG activity, were awakened from REM sleep to report dream mentation and to rate it on these variables. Coherence scores were calculated for homologous interhemispheric electrode pairs (Fp1-Fp2, F3-F4, F7-F8, C3-C4, P3-P4, O1-O2, T3-T4, T5-T6) and for left and right intrahemispheric pairs for delta, theta, alpha, beta1, and beta2 frequencies. These were correlated with the mentation measures. Positive correlations were found between average interhemispheric coherence in most bands and the character face measure. A breakdown by gender revealed that this relationship was most evident for women, whereas for men positive correlations were observed between coherence and negative self-feeling. That similar relationships also obtained for both left and right intrahemispheric coherence is consistent with the hypothesis that dreamed socioemotional interactions reflect the integrative functioning of many brain regions in both hemispheres.     

Differential responding to the beta movement after waking from REM and nonREM sleep.

Ten young adults were wakened from REM sleep and from nonREM sleep on two nonconsecutive nights and were tested to determine their upper and lower beta-movement thresholds. The ranges of the illusion were found to be significantly wider after waking from REM sleep than after waking from nonREM sleep or before sleep. The differential responding to the beta movement supports the experimental hypothesis that apparent motion may provide sensitive detectors of the operation during wakefulness of the Basic Rest-Activity Cycle, of which REM and nonREM sleep are opposite phases that carry over into wakefulness.     

Cortical arousal and mentation in sleeping and waking subjects

Cognitive variables and cortical arousal levels were examined in order to determine whether differences in cortical arousal levels within REM and waking could account for different aspects of mentation derived from the two states. Cognitive variables were derived from mentation reports collected from 30 subjects in both the waking state and after being awakened from REM sleep. Mentation reports were independently scored on seven content rating scales, by two judges blind to the conditions. These scales include among others, Total Recall Count (TRC), a count of all words in which the subject described his/her experience during the previous interval. The EEG activity, obtained from left and right midtemporoparietal and central sites, was recorded for 5-min periods before obtaining mentation reports. The absolute power of the EEG activity was calculated for each of six bandwidths. EEG power data from both waking and REM were entered into multiple linear regression equations to predict TRC. No relationships were found between TRC and general cortical activation, as measured by the EEG. Other statistical analyses, including relationships between EEG and scales of visual imagery, are discussed.     

Brain Gene Expression During REM Sleep Depends on Prior Waking Experience

In most mammalian species studied, two distinct and successive phases of sleep, slow wave (SW), and rapid eye movement (REM), can be recognized on the basis of their EEG profiles and associated behaviors. Both phases have been implicated in the offline sensorimotor processing of daytime events, but the molecular mechanisms remain elusive. We studied brain expression of the plasticity-associated immediate-early gene (IEG) zif-268 during SW and REM sleep in rats exposed to rich sensorimotor experience in the preceding waking period. Whereas nonexposed controls show generalized zif-268 down-regulation during SW and REM sleep, zif-268 is upregulated during REM sleep in the cerebral cortex and the hippocampus of exposed animals. We suggest that this phenomenon represents a window of increased neuronal plasticity during REM sleep that follows enriched waking experience.     

Cerebral blood flow in normal and abnormal sleep and dreaming

Measurements of regional or local cerebral blood flow (CBF) by the xenon-133 inhalation method and stable xenon computerized tomography CBF (CTCBF) method were made during relaxed wakefulness and different stages of REM and non-REM sleep in normal age-matched volunteers, narcoleptics, and sleep apneics. In the awake state, CBF values were reduced in both narcoleptics and sleep apneics in the brainstem and cerebellar regions. During sleep onset, whether REM or stage I-II, CBF values were paradoxically increased in narcoleptics but decreased severely in sleep apneics, while in normal volunteers they became diffusely but more moderately decreased. In REM sleep and dreaming CBF values greatly increased, particularly in right temporo-parietal regions in subjects experiencing both visual and auditory dreaming.     

Electroencephalographic intercentral interaction as a reflection of normal and pathological human brain activity

The authors summarized EEG findings and defined the nature of the intercentral EEG relationships in different functional states in healthy subjects and patients with organic cerebral pathology, based on a coherence analysis. Similar EEG characteristics in healthy individuals were identified: an anterior-posterior gradient of average coherence levels, the type of cortical-subcortical relationships in anterior cerebral structures. Right- and left-handed individuals showed frequent and regional differences in EEG coherence, which mainly reflected specificity of intracortical relationships. Development and regression of pathology in right-and left-handed individuals with organic brain lesions were thought to be caused by these differences. Lesions of regulatory structures (diencephalic, brain stem and limbic structures) provoked a more diffused kind of changes of intercentral relationships, in contrast to cortical pathology. These changes tended to reciprocate. The dynamic nature of intercentral relationships and their interhemispheric differences was revealed when changing functional states of the brain (increase and decrease of functional level) in healthy individuals and patients with organic cerebral pathology in the process of conscious and psychic activity restoration. Changing activity predominance of certain regulatory structures was considered one of the most important factors determining the dynamic nature of EEG coherence.     

Sleep and arousal mechanisms in experimental epilepsy: epileptic components of NREM and antiepileptic components of REM sleep

Neural generators related to different sleep components have different effects on seizure discharge. These sleep-related systems can provoke seizure discharge propagation during nonrapid eye movement (NREM) sleep and can suppress propagation during REM sleep. Experimental manipulations of discrete physiological components were conducted in feline epilepsy models, mostly in the systemic penicillin epilepsy model of primary generalized epilepsy and the amygdala kindling model of the localization-related seizure disorder, temporal lobe epilepsy. The sleep-wake state distribution of seizures was quantified before and after discrete lesions, systemic and localized drug administration, and/or photic stimulation, as well as in relation to microdialysis of norepinephrine. We found that (1) neural generators of synchronous EEG oscillations--including tonic background slow waves and phasic "arousal" events (sleep EEG transients such as sleep spindles and k-complexes)--combine to promote electrographic seizure propagation during NREM and drowsiness, and antigravity muscle tone permits seizure-related movement; (2) neural generators of asynchronous neuronal discharge patterns reduce electrographic seizures during alert waking and REM sleep, and skeletal motor paralysis blocks seizure-related movement during REM; (3) there are a number of similarities between amygdala-kindled kittens and children with Landau-Kleffner Syndrome (LKS) that suggest a link among seizures, sleep disorders, and behavioral abnormalities/regression.     

Electroencephalographic and autonomic alterations in subjects with frequent nightmares during pre-and post-REM periods

Abnormal arousal processes, sympathetic influences, as well as wake-like alpha activity during sleep were reported as pathophysiological features of Nightmare Disorder. We hypothesized that in Nightmare Disorder, wake-like cortical activity and peripheral measures linked to arousals would be triggered by physiological processes related to the initiation of REM periods. Therefore, we examined electroencephalographic (EEG), motor and autonomous (cardiac) activity in a group of nightmare (NM) and healthy control (CTL) subjects during sleep-state-transitions while controlling for the confounding effects of trait anxiety. Based on the second-nights’ polysomnographic recordings of 19 Nightmare Disordered (NM) and 21 control (CTL) subjects, we examined the absolute power spectra focusing on the alpha range, measures of heart rate variability (HRV) and motor (muscle tone) activity during pre-REM and post-REM periods, separately. According to our results, the NM group exhibited increased alpha power during pre-REM, but not in post-REM, or stable, non-transitory periods. While CTL subjects showed increased HRV during pre-REM periods in contrast to post-REM ones, NM subjects did not exhibit such sleep state-specific differences in HRV, but showed more stable values across the examined sleep stages and less overall variability reflecting generally attenuated parasympathetic activity during sleep-state-transitions and during stable, non-transitory NREM states. These differences were not mediated by waking levels of trait anxiety. Moreover, in both groups, significant differences emerged regarding cortical and motor (muscle tone) activity between pre-REM and post-REM conditions, reflecting the heterogeneity of NREM sleep. Our findings indicate that NM subjects’ sleep is compromised during NREM–REM transitions, but relatively stabilized after REM periods. The coexistence of sleep-like and wake-like cortical activity in NM subjects seems to be triggered by REM/WAKE promoting neural activity. We propose that increased arousal-related phenomena in NREM–REM transitions might reflect altered emotional processing in NM subjects.     

Bimanual coordination and interhemispheric interaction

Bimanual coordination of skilled finger movements requires intense functional coupling of the motor areas of both cerebral hemispheres. This coupling can be measured non-invasively in humans with task-related coherence analysis of multi-channel surface electroencephalography. Since bimanual coordination is a high-level capability that virtually always requires training, this review is focused on changes of interhemispheric coupling associated with different stages of bimanual learning. Evidence is provided that the interaction between hemispheres is of particular importance in the early phase of command integration during acquisition of a novel bimanual task. It is proposed that the dynamic changes in interhemispheric interaction reflect the establishment of efficient bimanual ‘motor routines'. The effects of callosal damage on bimanual coordination and learning are reviewed as well as functional imaging studies related to bimanual movement. There is evidence for an extended cortical network involved in bimanual motor activities which comprises the bilateral primary sensorimotor cortex (SM1), supplementary motor area, cingulate motor area, dorsal premotor cortex and posterior parietal cortex. Current concepts about the functions of these structures in bimanual motor behavior are reviewed.     

Coherence of hemispheric function in developmental dyslexia

Left parieto-occipital EEG leads record a frequency spectrum in dyslexics that is consistently different from the spectrum obtained from normals. It is suggested that these effects represent significant differences in the functional organization of these areas. EEG coherence values indicate that normals have significantly greater sharing between hemispheres at symmetrical locations. Dyslexics demonstrate significantly greater sharing within hemisphere than do normals. The data supports developmental dyslexia being a functional hemispheric disconnection syndrome.     

Motivation and affect in REM sleep and the mentation reporting process

Although the emotional and motivational characteristics of dreaming have figured prominently in folk and psychoanalytic conceptions of dream production, emotions have rarely been systematically studied, and motivation, never. Because emotions during sleep lack the somatic components of waking emotions, and they change as the sleeper awakens, their properties are difficult to assess. Recent evidence of limbic system activation during REM sleep suggests a basis in brain architecture for the interaction of motivational and cognitive properties in dreaming. Motivational and emotional content in REM and NREM laboratory mentation reports from 25 participants were compared. Motivational and emotional content was significantly greater in REM than NREM sleep, even after controlling for the greater word count of REM reports.     

Dreaming and the brain: toward a cognitive neuroscience of conscious states

Sleep researchers in different disciplines disagree about how fully dreaming can be explained in terms of brain physiology. Debate has focused on whether REM sleep dreaming is qualitatively different from nonREM (NREM) sleep and waking. A review of psychophysiological studies shows clear quantitative differences between REM and NREM mentation and between REM and waking mentation. Recent neuroimaging and neurophysiological studies also differentiate REM, NREM, and waking in features with phenomenological implications. Both evidence and theory suggest that there are isomorphisms between the phenomenology and the physiology of dreams. We present a three-dimensional model with specific examples from normally and abnormally changing conscious states.     

The role of sleep in changing our minds: a psychologist's discussion of papers on memory reactivation and consolidation in sleep

The group of papers on memory reactivation and consolidation during sleep included in this volume represents cutting edge work in both animals and humans. They support that the two types of sleep serve different necessary functions. The role of slow wave sleep (SWS) is reactivation of the hippocampal-neocortical circuits activated during a waking learning period, while REM sleep is responsible for the consolidation of this new learning into long-term memory. These studies provide further insights into mechanisms involved in brain plasticity. Robeiro has demonstrated the upregulation of an immediate-early gene (IEG zif 268) to waking levels, which occurs only in REM and only in connection with new learning. McNaughton and his group have identified electrical indicators that the hippocampus and neocortex are talking to each other by testing the coactivation of hippocampal sharp wave bursts in SWS and shifts from down to up states of activation in the neocortex. In human studies Smith's group reports work on individual differences such as intelligence and presleep alcohol that affect postsleep performance, and Stickgold and collaborators report that a short nap will improve performance if it contains REM sleep. Payne and Nadel suggest that the recall benefit associated with REM sleep may be due to its association with increased cortisol levels. These papers are important not only in their individual contributions but also in revitalizing the work coordinating waking and sleep. This promises to further the understanding of how our unique capacity to learn from experience and modify our behavior takes place.     

Search activity in the context of psychosomatic disturbances,of brain monoamines and REM sleep function

The author discusses a number of controversial aspects of the search activity concept. This concept, based on an analysis of data cited by other researchers and the results of the author’s own investigation, performed together with V. V. Arshavsky, postulates that search activity raises the body’s resistance to stress and experimentally induced pathology whereas renunciation of search forms a nonspecific predisposition to somatic disturbances (e.g., psychosomatic disease). REM sleep is regarded as a specific form of search activity aimed at compensating for the state of renunciation of search in walking. In this paper the author argues that 1) renunciation of search can be accompanied either by anxiety or by depression, 2) REM sleep deprivation on a “small platform” raises the requirement in REM sleep by producing renunciation of search, 3) during search activity brain catecholamine synthesis is stimulated by catabolism whereas a state of renunciation of search upsets this feedback system. The actuation of the brain mechanisms of search in REM sleep necessitates a certain brain catecholamine level. If the brain catecholamine level is very high during waking behavior due to intensive search activity, the REM sleep requirement is low, REM sleep becoming reduced. After a moderate drop in the brain catecholamine level at the initial stage of renunciation of search the requirement in REM sleep rises and this phase grows longer. But at the late stage of renunciation of search the brain catecholamine level drops extremely, REM sleep shrinking in spite of the great appropriate requirement, and 4) the functional insufficiency of REM sleep invites various forms of pathology.     

Dreaming and the default network: A review,synthesis, and counterintuitive research proposal

This article argues that the default network, augmented by secondary visual and sensorimotor cortices, is the likely neural correlate of dreaming. This hypothesis is based on a synthesis of work on dream content, the findings on the contents and neural correlates of mind-wandering, and the results from EEG and neuroimaging studies of REM sleep. Relying on studies in the 1970s that serendipitously discovered episodes of dreaming during waking mind-wandering, this article presents the seemingly counterintuitive hypothesis that the neural correlates for dreaming could be further specified in the process of carrying out EEG/fMRI studies of mind-wandering and default network activity. This hypothesis could be tested by asking participants for experiential reports during moments of differentially high levels of default network activation, as indicated by mixed EEG/fMRI criteria. Evidence from earlier EEG/fMRI studies of mind-wandering and from laboratory studies of dreaming during the sleep-onset process is used to support the argument.     

Behavioral,subjective, and electroencephalographic indices of sleep onset latency and sleep duration

Electrographic (EEG, EOG, EMG) indices have been used for some decades in the definition of the stages of sleep and more recently in the diagnosis of sleep-related disorders, e.g., insomnia, despite the lack of detailed information concerning the precise relationship between such electrographic indices and behavioral and subjective criteria of sleep. Evaluation of the relationship between EEG, behavioral, and self-report measures of sleep onset latency and sleep duration was conducted using 17 young normal sleepers. The behavioral measure was provided by an apparatus which records subjects' button-press responses to an auditory stimulus presented at various interstimulus intervals (2, 5, and random 1, 2, 5, 7, and 10 min). The behavioral and stage 2 EEG estimates of sleep onset latency (SOL) and sleep duration (SD) were almost identical. The stage 1 EEG provided the shortest estimate of SOL; the self-report measure, the longest. The SD measures were in reverse order. There were no significant differences among the three interstimulus interval conditions (2, 5, and random min). Average response rates to the 50-dB chime were 100% during wakefulness, 81% during EEG stage 1, and 8% during EEG stage 2. Almost all stage 2 responding occurred during the first 5 min of each stage 2 period. The validity of electrographic indices as sleep criteria and the implications of the findings for the formulation of an adequate definition of sleep and its clinical measurement are discussed.     

Slow wave sleep and recollection in recognition memory

Recognition memory performance reflects two distinct memory processes: a conscious process of recollection, which allows remembering specific details of a previous event, and familiarity, which emerges in the absence of any conscious information about the context in which the event occurred. Slow wave sleep (SWS) and rapid eye movement (REM) sleep are differentially involved in the consolidation of different types of memory. The study assessed the effects of SWS and REM sleep on recollection, by means of the "remember"/"know" paradigm. Subjects studied three blocks of 12 words before a 3-h retention interval filled with SWS, REM sleep or wakefulness, placed between 3 a.m. and 6 a.m. Afterwards, recognition and recollection were tested. Recollection was higher after a retention interval rich in SWS than after a retention interval rich in REM sleep or filled with wakefulness. The results suggest that SWS facilitates the process of recollection in recognition memory.     

REM periodicity under ultrashort sleep/wake cycle in narcoleptic patients

Six narcoleptic patients were tested three times on the 13-min waking/7-min resisting sleep paradigm each time after a night of sleep in the laboratory. The three experiments were conducted after 10 days without any antinarcoleptic treatment or after 2 weeks of daily treatment with either methyl-phenidate or aniracetam. The results showed that patients had pronounced levels of diurnal sleepiness in all three experimental conditions with a midafternoon peak at around 1300-1500 hr and a nadir at around 1800 hr. Methyl-phenidate significantly reduced REM sleep and marginally reduced total sleep in comparison with the no-treatment and aniracetam conditions. REM sleep in the 7/13 paradigm appeared cyclically with a dominant periodicity of 80 min/cycle. The cycles tended to be synchronized across patients and were unrelated to the temporal structure of total sleep. The present results support the continuation of the REM oscillator during brief periods of waking, but suggest that the REM periodicity is unrelated to Kleitman's BRAC model of arousal.