Dynamics of frontal EEG activity, sleepiness and body temperature under high and low sleep pressure.

The impact of sleep deprivation (high sleep pressure) vs sleep satiation (low sleep pressure) on waking EEG dynamics, subjective sleepiness and core body temperature (CBT) was investigated in 10 young volunteers in a 40 h controlled constant posture protocol. The differential sleep pressure induced frequency-specific changes in the waking EEG from 1-7 Hz and 21-25 Hz. Frontal low EEG activity (FLA, 1-7 Hz) during sleep deprivation exhibited a prominent increase as time awake progressed, which could be significantly attenuated by sleep satiation attained with intermittent naps. Subjective sleepiness exhibited a prominent circadian regulation during sleep satiation, with virtually no homeostatic modulation. These extremely different sleep pressure conditions were not reflected in significant changes of the CBT rhythm. The data demonstrate that changes in FLA during wakefulness are to a large extent determined by the sleep-wake dependent process with little circadian modulation, and reflect differential levels of sleep pressure in the awake subject.     

High frequency waking EEG: reflection of a slow ultradian rhythm in daytime arousal

The ultradian dynamics of the human waking EEG was studied using a short visual fixation task repeated every 10 min throughout the daytime. The EEG spectra obtained from the tasks were assessed for time effect and ultradian periodicity. Fronto-central EEG high frequency powers (22.5-44.5 Hz) decreased at the time of the midafternoon vigilance dip (14.00-17.00 h) along with slight concomitant increases in parietal alpha (7.5-13.5 Hz) and delta (1-3 Hz) powers. A slow ultradian rhythm with a 3-4 h periodicity strongly modulated EEG power in all frequency bands between 1 and 44.5 Hz. The high frequency waking EEG may well reflect the activity of a brain arousal process underlying maintenance of the waking state probably throughout the 24 h cycle.     

Spectral analysis of slow modulation of EEG amplitude and cardiovascular variables in subjects with postural tachycardia syndrome

OBJECTIVE: Previous studies have reported slow (<0.5 Hz) modulation of electroencephalographic (EEG) background amplitude and suggested that this reflects periodic neuronal activity in the brainstem, such as may be recorded from cardiovascular and respiratory centers in animals. We searched for a relationship between EEG amplitude modulation and modulation of simultaneously recorded cardiovascular variables and attempted to determine whether this relationship was altered in subjects with postural tachycardia syndrome (POTS). METHODS: We recorded EEG, blood flow velocity in the middle cerebral artery (MCA), heart rate, respirations, and blood pressure from subjects with POTS and controls during head-up tilt. Time-frequency analysis of 0.512-s epochs of EEG was performed to determine peak alpha amplitude. Spectra were divided into 3 bands: ultraslow, middle, and respiratory. RESULTS: EEG alpha amplitude modulation in all frequency bands was reduced in POTS subjects while supine. EEG modulation decreased in controls with head-up tilt but not in POTS subjects. Heart rate modulation in the respiratory frequency band decreased with head-up tilt and was significantly less (P<0.02) in ultraslow and respiratory frequency bands in POTS subjects after head-up tilt. Blood pressure and MCA flow velocity modulation in middle and respiratory bands increased with head-up tilt to a greater degree in POTS subjects. Blood pressure and MCA flow velocity modulation frequencies were moderately correlated, but correlations between EEG and cardiovascular variable modulation frequencies were generally low, being highest in the respiratory band but not statistically significant. CONCLUSION: There are subtle differences in EEG amplitude modulation in subjects with POTS. Altered EEG amplitude modulation in POTS may reflect altered brainstem physiology in this disorder.     

Blood pressure and heart rate variability in autonomic disorders: a critical review

Spectral analysis (SA) of blood pressure (BP) and heart rate (HR) fluctuations has been proposed as a unique approach to obtain a deeper insight into cardiovascular regulatory mechanisms in health and disease. A number of studies performed over the last 15 years have shown that autonomic influences are involved in the modulation of fast BP and HR fluctuations (with a period<1 min), particularly at frequencies between 0.2 and 0.4 Hz [high frequency (HF) region or respiratory frequency] and around 0.1 Hz [mid frequency (MF) region]. In patients with secondary or primary autonomic dysfunction, SA of BP and HR signals recorded at rest or during orthostatic challenge in a laboratory environment have shown the occurrence of a reduction in the power of MF and/or HF, BP and HR components. Such a reduction is associated or may even precede the clinical manifestation of autonomic neuropathy. However, the above results collected in standardized laboratory conditions cannot reflect the features of neural cardiovascular control during daily life in ambulant individuals with autonomic failure. To investigate this issue, SA techniques have been applied to 24 h beat-to-beat intra-arterial and non-invasive finger BP recordings obtained in elderly subjects and in pure autonomic failure patients, respectively. In these conditions, HR powers displayed a reduction over a wide range of frequencies (from 0.5 to below 0.01 Hz). Conversely, BP powers underwent a complex rearrangement characterized by a reduction in the power around 0.1 Hz and by an increase in the powers at the respiratory frequency and at frequencies below 0.01 Hz. Dynamic quantification of the sensitivity of the baroreceptor-heart rate reflex by combined analysis of systolic BP and pulse interval (i.e. the interval between consecutive systolic peaks) powers around 0.1 Hz (alpha technique) has shown that in elderly subjects, and even more so in pure autonomic failure patients, baroreflex sensitivity is markedly reduced over the 24 h, and is no longer characterized by its physiological day-night modulation. In conclusion, although in some instances SA of cardiovascular signals may fail to fully reflect the features of autonomic cardiovascular control, the evidence discussed clearly demonstrates that this approach represents a promising tool for a dynamic assessment of the early impairment of neural circulatory control in autonomic failure. This is particularly the case when these analyses are performed on 24 h continuous BP and HR recordings in ambulant subjects.     

Potentials evoked by chirp-modulated tones: a new technique to evaluate oscillatory activity in the auditory pathway.

OBJECTIVE: Steady-state potentials are oscillatory responses generated by a rhythmic stimulation of a sensory pathway. The frequency of the response, which follows the frequency of stimulation, is maximal at a stimulus rate of 40 Hz for auditory stimuli. The exact cause of these maximal responses is not known, although some authors have suggested that they might be related to the 'working frequency' of the auditory cortex. Testing of the responses to different frequencies of stimulation may be lengthy if a single frequency is studied at a time. Our aim was to develop a fast technique to explore the oscillatory response to auditory stimuli, using a tone modulated in amplitude by a sinusoid whose frequency increases linearly in frequency ('chirp') from 1 to 120 Hz. METHODS: Time-frequency transforms were used for the analysis of the evoked responses in 10 subjects. Also, we analyzed whether the peaks in these responses were due to increases of amplitude or to phase-locking phenomena, using single-sweep time-frequency transforms and inter-trial phase analysis. RESULTS: The pattern observed in the time-frequency transform of the chirp-evoked potential was very similar in all subjects: a diagonal band of energy was observed, corresponding to the frequency of modulation at each time instant. Two components were present in the band, one around 45 Hz (30-60 Hz) and a smaller one between 80 and 120 Hz. Inter-trial phase analysis showed that these components were mainly due to phase locking phenomena. CONCLUSIONS: A simultaneous testing of the amplitude-modulation-following oscillatory responses to auditory stimulation is feasible using a tone modulated in amplitude at increasing frequencies. The maximal energies found at stimulation frequencies around 40 Hz are probably due to increased phase-locking of the individual responses.     

The mechanics of pigeon eye movements--are they like other vertebrates?

The lateral rectus muscle of the pigeon eye was driven by stimulation of the abducens nucleus. Eye rotation was measured with an opto-electronic movement detector. Eye position was linearly related to stimulation frequency in the range 40-110 Hz and saturated at frequencies above 250 Hz. Maximum eye velocity of 240 degrees/s was obtained with a stimulation frequency of 360 Hz. Stimulation with sinusoidally modulated pulse frequencies (40-110 Hz) over the modulation frequency range 0.01-6.0 Hz were used to determine the gain and phase relationships of the oculomotor system. The response approximates a first order low pass frequency model with a characteristic frequency of 0.45 Hz at high frequencies. There is an additional phase lag equivalent to a time delay of 9.8 ms. The results are compared with similar experiments performed on the dogfish and cat oculomotor systems.     

Neurofeedback in healthy elderly human subjects with electroencephalographic risk for cognitive disorder

In normal elderly subjects, the best electroencephalogram (EEG)-based predictor of cognitive impairment is theta EEG activity abnormally high for their age. The goal of this work was to explore the effectiveness of a neurofeedback (NFB) protocol in reducing theta EEG activity in normal elderly subjects who present abnormally high theta absolute power (AP). Fourteen subjects were randomly assigned to either the experimental group or the control group; the experimental group received a reward (tone of 1000 Hz) when the theta AP was reduced, and the control group received a placebo treatment, a random administration of the same tone. The results show that the experimental group exhibits greater improvement in EEG and behavioral measures. However, subjects of the control group also show improved EEG values and in memory, which may be attributed to a placebo effect. However, the effect of the NFB treatment was clear in the EG, although a placebo effect may also have been present.     

Regional differences in the circadian modulation of human sleep spindle characteristics

Electroencephalographic oscillations in the sleep spindle frequency range (11-16 Hz) are a key element of human nonrapid eye movement sleep. In the present study, sleep spindle characteristics along the anterior-posterior axis were analysed during and outside the circadian phase of melatonin secretion. Sleep electroencephalograms were recorded during naps distributed over the entire circadian cycle and analysed with two different methodological approaches, the classical fast Fourier transform in the frequency-domain and a new method for instantaneous spectral analysis, the fast time frequency transform that yields high-resolution parameters in the combined time-frequency-domain. During the phase of melatonin secretion, spindle density was generally increased and intraspindle frequency variation reduced. Furthermore, lower spindle frequencies were promoted: peak frequencies shifted towards the lower end of the spindle frequency range, and spindle amplitude was enhanced in the low-frequency range (11-14.25 Hz) and reduced in the high-frequency range (approximately 14.5-16 Hz). The circadian variation showed a clear dependence on brain topography such that it was maximal in the parietal and minimal in the frontal derivation. Our data provide evidence that the circadian pacemaker actively promotes low-frequency sleep spindles during the biological night with a parietal predominance.     

EEG spectral power and sleepiness during 24 h of sustained wakefulness in patients with obstructive sleep apnea syndrome.

OBJECTIVE: This study investigated if obstructive sleep apnea syndrome (OSAS) may be associated with higher activity in different frequency bands of the EEG during a sustained wakefulness paradigm. METHODS: Twelve OSA patients and 8 healthy controls were studied with the Karolinska Drowsiness Test (KDT) and subjective ratings of sleepiness (VAS and KSS) conducted every hour during 24 h of sustained wakefulness. RESULTS: The waking EEG activity, mainly in the low (0.5-7.8 Hz) and fast (12.7-29.2 Hz) frequency band, increased as time awake progressed in both groups but more obviously in OSA patients. A similar pattern was observed for rated sleepiness in both groups. Moreover, VAS ratings of alertness were closely related to the awake theta, fast alpha and beta bands in controls but not in OSA patients. CONCLUSIONS: OSAS was associated with a wake-dependent increase in low (0.5-7.8 Hz) and fast (12.7-29.2 Hz) frequency range activity. Variations in behavioural sleepiness measured by VAS ratings closely reflect most of the waking EEG parameters in controls but not in OSA patients. SIGNIFICANCE: In a sustained wakefulness paradigm, higher activity in delta, theta and beta bands associated with OSAS indicates that OSA patients show marked signs of higher sleepiness and stronger efforts than controls to stay awake, even though they tend to underestimate their sleepiness.     

Human VEP contrast modulation sensitivity: separation of magno- and parvocellular components.

Human cortical visual evoked potentials (VEPs) were retrieved in real time (without averaging). The stimuli were sinusoidal gratings whose contrast was temporally modulated about some mean value. Electrophysiologically determined contrast modulation thresholds were measured at standing contrast over the range from 2.5% to 50%, defining an increment threshold function. Increment threshold functions were obtained under two different spatio-temporal stimulus conditions identified as "sustained" (4 c/deg grating modulated at 1.5 Hz) and "transient" (1 c/deg grating modulated at 20 Hz). Under each of these conditions, threshold responses were retrieved at both the fundamental and second harmonic of the contrast modulation frequency. Under "sustained" conditions log increment threshold responses and the fundamental the second harmonic of the modulation frequency were similar to those at the fundamental except for a saturation effect (i.e., above a mean contrast of 25% there was little further reduction in modulation sensitivity). There was no contrast gain control under "transient" stimulus conditions. In other words, the same absolute amount of contrast change produced threshold responses for all mean levels up to 25%. This was true at both the fundamental and second harmonic of the modulation frequency. Stimulus differences produce striking differences in the electrophysiologically inferred increment threshold function for grating contrast, but fundamental and second harmonic evoked responses reflect processes with similar increment threshold behavior.     

Functional topography of the human nonREM sleep electroencephalogram

The sleep EEG of healthy young men was recorded during baseline and recovery sleep after 40 h of waking. To analyse the EEG topography, power spectra were computed from 27 derivations. Mean power maps of the nonREM sleep EEG were calculated for 1-Hz bins between 1.0 and 24.75 Hz. Cluster analysis revealed a topographic segregation into distinct frequency bands which were similar for baseline and recovery sleep, and corresponded closely to the traditional frequency bands. Hallmarks of the power maps were the frontal predominance in the delta and alpha band, the occipital predominance in the theta band, and the sharply delineated vertex maximum in the sigma band. The effect of sleep deprivation on EEG topography was determined by calculating the recovery/baseline ratio of the power spectra. Prolonged waking induced an increase in power in the low-frequency range (1-10.75 Hz) which was largest over the frontal region, and a decrease in power in the sigma band (13-15.75 Hz) which was most pronounced over the vertex. The topographic pattern of the recovery/baseline power ratio was similar to the power ratio between the first and second half of the baseline night. These results indicate that changes in sleep propensity are reflected by specific regional differences in EEG power. The predominant increase of low-frequency power in frontal areas may be due to a high 'recovery need' of the frontal heteromodal association areas of the cortex.     

Perceptual learning evidence for tuning to spectrotemporal modulation in the human auditory system

Natural sounds are characterized by complex patterns of sound intensity distributed across both frequency (spectral modulation) and time (temporal modulation). Perception of these patterns has been proposed to depend on a bank of modulation filters, each tuned to a unique combination of a spectral and a temporal modulation frequency. There is considerable physiological evidence for such combined spectrotemporal tuning. However, direct behavioral evidence is lacking. Here we examined the processing of spectrotemporal modulation behaviorally using a perceptual-learning paradigm. We trained human listeners for ～1 h/d for 7 d to discriminate the depth of spectral (0.5 cyc/oct; 0 Hz), temporal (0 cyc/oct; 32 Hz), or upward spectrotemporal (0.5 cyc/oct; 32 Hz) modulation. Each trained group learned more on their respective trained condition than did controls who received no training. Critically, this depth-discrimination learning did not generalize to the trained stimuli of the other groups or to downward spectrotemporal (0.5 cyc/oct; -32 Hz) modulation. Learning on discrimination also led to worsening on modulation detection, but only when the same spectrotemporal modulation was used for both tasks. Thus, these influences of training were specific to the trained combination of spectral and temporal modulation frequencies, even when the trained and untrained stimuli had one modulation frequency in common. This specificity indicates that training modified circuitry that had combined spectrotemporal tuning, and therefore that circuits with such tuning can influence perception. These results are consistent with the possibility that the auditory system analyzes sounds through filters tuned to combined spectrotemporal modulation.     

EEG and subjective sleepiness during extended wakefulness in seasonal affective disorder: circadian and homeostatic influences.

BACKGROUND: Seasonal affective disorder (SAD) may reflect a disturbance of circadian phase relationships or a disturbance of sleep-wake dependent processes, both of which change daytime energy and sleepiness levels. METHODS: Under the unmasking conditions of a 40-hour constant routine protocol (CR), self-rated sleepiness and waking electroencephalogram (EEG) power density were assessed in women with SAD (n = 8) and in age-matched healthy control subjects (n = 9). RESULTS: There was no significant effect of season or light treatment in any of the measures.The time course of subjective sleepiness was characterized by a circadian modulation and an overall increase during extended wakefulness in both SAD patients and control subjects. A prominent circadian rhythm of subjective sleepiness was not different in SAD patients and control subjects; however, the progressive buildup of sleepiness, as quantified by nonlinear regression analysis, was significantly reduced in SAD patients, mainly because they were sleepier than control subjects during the first 12 hours of the CR. The time course of waking EEG theta-alpha activity showed a more rapid increase during the first 10 hours of the CR in SAD patients. In contrast to control subjects who showed a progressive increase in the course of the 40-hour episode of extended wakefulness, EEG theta-alpha activity in SAD patients did not further increase over the remainder of the CR. CONCLUSIONS: The data suggest that SAD patients may have a trait (rather than state) deficiency in the homeostatic buildup of sleep pressure during extended wakefulness as indexed by subjective sleepiness and EEG theta-alpha activity.     

Circadian periodicity of heart rate variability in hospitalized angor patients

Abstract The relationship between unstable angor (angina) and circadian periodicity of heart rate variability (HRV) was explored in a group of patients hospitalized in a coronary care unit (CCU). Patients were classified as normal (whose symptoms had non-cardiovascular origin, n=8), moderate angor (n=13) and severe angor (n=11). A fourth group of ambulatory healthy volunteers (n=12) was included. Individual 24 h Holter records were analyzed, mean RR and standard deviation of RR (SDNN) being obtained from 1 h-length windows. For frequency domain analysis, 5 min-length windows were employed. The spectral components analyzed were total power (spectral power between 0.01 and 0.5 Hz), low frequency power (LF: power between 0.04 and 0.15 Hz), and high frequency power (HF: power between 0.15 and 0.4 Hz). In addition, LF to HF areas ratio (L/H) was computed. Mesor, amplitude and acrophase for every 24 h rhythm were calculated by cosinor analysis.As compared to ambulatory controls, admission to the CCU diminished amplitude and phase-delayed the circadian oscillation of most HRV parameters, except for SDNN. Moderate angor patients showed decreased amplitude of RR and L/H and augmented amplitude of SDNN when compared to normal hospitalized subjects. A phase delay of about 1.5 h for RR intervals and a phase advance of 3.5–6 h for LFA and SDNN were found in the moderate angor group when compared to normal. Amplitude of 24 h variation of total power decreased in severely angor patients and the circadian oscillation of HF (an indicator of vagal control on the heart) became free running. A phase delay of 2.5 h in SDNN acrophase was found in severely affected patients when compared to moderate. The results indicate that severity of unstable angor correlates with desynchronization of parasympathetic control of heart rate.     

Spectral analysis of adult dancers' sways: sex and interaction vision-proprioception

In subjects of both sexes with or without dance training, dependence on vision and proprioception for postural control was studied by destabilizing these cues on a free seesaw. Fast Fourier transform processing allowed spectral frequency analysis of the platform sways recorded by an accelerometer. Two frequency bands of the total spectral energy were used: the lower (0 - 2 Hz) and the higher (2 - 20 Hz) frequency bands. Dancers were significantly less dependent on vision but use more proprioception than untrained subjects. Professional dance training appears to shift sensorimotor dominance from vision to proprioception, and this evolution seems more marked for males than females. Female and male dancers had similar dynamic performances, but for males, the better neuromuscular coordination may be associated with biomechanical factors.     

Effects of circadian phase and duration of sleep deprivation on sleep and EEG power spectra in the cat

The electroencephalogram (EEG) of cats was recorded under baseline conditions (LD 12:12) and after 4 and 8 h of sleep deprivation (SD). The EEG was analyzed by visual scoring and by spectral analysis. Under baseline conditions the 24-h distribution of sleep was bimodal: the smallest amounts of sleep occurred at the light-dark and dark-light transitions. EEG slow-wave activity (power density in the delta frequency range: 0.5-4.0 Hz) in non-rapid-eye-movement sleep (NREMS) showed a small variation over the 24-h period. When recovery sleep, following 4 h and 8 h of SD, started at the beginning of the dark period, no significant rebound of NREMS and REMS occurred during the 24-h recovery period. When recovery sleep, after 4 h of SD, started at the fifth hour of the light period, the amount of NREMS was increased. In all experiments the EEG power density in NREMS was enhanced after SD in the entire frequency range studied (0.5-31.5 Hz), but more prominently in the delta and theta (4.5-7.0 Hz) frequency bands. The effects dissipated in the course of the recovery period. The magnitude and duration of the enhancements of EEG power densities were dependent on the duration of SD and on the circadian phase at which SD was scheduled. It is concluded that in the cat sleep is a function of both circadian and homeostatic processes and that especially the EEG power density in NREMS is highly responsive to sleep loss.     

Developmental changes of distortion product and transient evoked otoacoustic emissions in different age groups

The developmental changes of distortion product otoacoustic emissions (DPOAEs) and transient evoked otoacoustic emissions (TEOAEs) were evaluated in 275 normal subjects aged from 1 month to 39 years. The DP-grams showed an M-shaped pattern with peaks at 1587 Hz and 5042 Hz for all age groups. In subjects younger than 3 years, low frequency DPOAEs did not rise above the noise floor. The DP levels at high frequency (5042 Hz) did not change much from infancy to young adulthood (12.9-16.5 dB SPL), however, those at low and middle frequency significantly decreased with age. Total echo power (TEP) of TEOAE was greatest in early infancy, decreased rapidly before 6-7 years old, and then decreased gradually (TEP = 16.6 - 1.9 X ln (age)). Wave reproducibility was constant across age. The frequency area peak power (FAPP) to middle and high frequency sounds changed little with age, however, FAPP at low frequency sounds dramatically increased with age. FAPP at 5000 Hz was relatively depressed levels at each age. The TEOAE value was more prominent at middle and low frequencies while DPOAE was predominant at high frequencies. These two measurements may reflect different functions of outer hair cells in the developing cochlea.     

Cortical sound processing in children with autism disorder: an MEG investigation

Previous work investigating frequency encoding mechanisms in human auditory cortex has provided evidence that latency of the auditory evoked M100 is strongly proportional to frequency, with low frequency (100-200 Hz) tones associated with approximately 30 ms longer latencies than mid-range frequency (1-2 kHz) tones. Motivated by pervasive speech and auditory perception deficits observed in autism spectrum disorder, we evaluated M100 frequency dependence in children with autism disorder and typically developing controls. Results indicate that for control children, the dynamic range of frequency modulation was similar to previous reports for healthy adults. Children with autism had a much reduced range of modulation in right hemisphere sites. Findings indicate that frequency encoding mechanisms may follow a differential maturational path in autism spectrum disorder.     

Variable otolith contribution to the galvanically induced vestibulo-ocular reflex.

The torsional eye movements elicited by sinusoidal galvanic vestibular stimulation (GVS) (0.012-3.13 Hz) were examined in healthy humans. GVS consistently induced sinusoidal modulation of the torsional slow phase velocity (SPV), which was linearly related to stimulus intensity. At low frequencies (< 0.1 Hz) nystagmic responses could be discriminated from an underlying 'tonic' modulation of eye position, which was prominent in some, but negligible in other subjects, and was not correlated with the SPV modulation. The actual SPV modulation consistently exceeded the (hypothetical) velocity modulation derived from the tonic positional components, albeit variably by almost 20-fold across subjects. This indicates that the contribution of possibly otolith-related response components to the galvanic vestibulo-ocular reflex may vary considerably in normal individuals.     

Endogenously generated gamma‐band oscillations in early visual cortex: A neurofeedback study

Human subjects were trained with neurofeedback (NFB) to enhance the power of narrow‐band gamma oscillations in circumscribed regions of early visual cortex. To select the region and the oscillation frequency for NFB training, gamma oscillations were induced with locally presented drifting gratings. The source and frequency of these induced oscillations were determined using beamforming methods. During NFB training the power of narrow band gamma oscillations was continuously extracted from this source with online beamforming and converted into the pitch of a tone signal. We found that seven out of ten subjects were able to selectively increase the amplitude of gamma oscillations in the absence of visual stimulation. One subject however failed completely and two subjects succeeded to manipulate the feedback signal by contraction of muscles. In all subjects the attempts to enhance visual gamma oscillations were associated with an increase of beta oscillations over precentral/frontal regions. Only successful subjects exhibited an additional marked increase of theta oscillations over precentral/prefrontal and temporal regions whereas unsuccessful subjects showed an increase of alpha band oscillations over occipital regions. We argue that spatially confined networks in early visual cortex can be entrained to engage in narrow band gamma oscillations not only by visual stimuli but also by top down signals. We interpret the concomitant increase in beta oscillations as indication for an engagement of the fronto‐parietal attention network and the increase of theta oscillations as a correlate of imagery. Our finding support the application of NFB in disease conditions associated with impaired gamma synchronization.