Fronto-parietal alpha ERD and visuo-spatial attention in pregnant women.

Many pregnant women report impairments in their attentional capacities. However, comparative studies between pregnant and non-pregnant women using standardised attention paradigms are rare and inconsistent. During attention tasks alpha activity is known to suppress irrelevant sensory inputs and previous studies show that a large event-related desynchronisation (ERD) in the alpha range prior to target-onset predicts enhanced attentional processing. We quantified the relationship between performance (accuracy, response time) in a standardised visuo-spatial attention task and alpha ERD (∼6-12 Hz) as well as saliva estradiol level in fifteen pregnant women (M = 26.6, SD = 3.0 years) compared to fifteen non-pregnant, naturally cycling women (M = 23.1, SD = 4.3 years). Compared to non-pregnant women, alpha frequency was increased in pregnant women. Furthermore, pregnant women showed a greater magnitude of the alpha ERD prior to target-onset and a moderate increase in accuracy compared to non-pregnant women. In addition, accuracy correlated negatively with estradiol in pregnant women as well as with frontal alpha ERD in all women. These correlational findings indicate that pregnancy-related enhancement in alpha desynchronisation in a fronto-parietal network might modulate accuracy during a visuo-spatial attention task.

The Pavlovian craver: Neural and experiential correlates of single trial naturalistic food conditioning in humans.

Present-day environments are replete with tempting foods and the current obesity pandemic speaks to humans' inability to adjust to this. Pavlovian processes may be fundamental to such hedonic overeating. However, a lack of naturalistic Pavlovian paradigms in humans makes translational research difficult and important parameters such as implicitness and acquisition speed are unknown. Here we present a novel naturalistic conditioning task: an image of a neutral object was conditioned to marzipan taste in a single trial procedure by asking the participant to eat the 'object' (made from marzipan). Relative to control objects, results demonstrate robust pre- to post-conditioning changes of both subjective ratings and early as well as late event related brain potentials, suggesting contributions of implicit (attentional) and explicit (motivational) processes. Naturalistic single-trial taste-appetitive conditioning is potent in humans and shapes attentional and motivational neural processes that might challenge self-regulation during exposure to tempting foods. Thus, appetitive conditioning processes might contribute to overweight and obesity.

Associations of endogenous 17-ß-estradiol with theta amplitude and performance in semantic categorization in young women.

In semantic categorization processes, individuals form a relation between perceived or imagined objects and knowledge about these objects. In the present semantic categorization study, we correlated endogenous 17-ß-estradiol levels (E2) with performance as well as amplitude of theta oscillations in young women (age 23.1±3.4 years). The semantic categorization task consisted of nouns representing either living or non-living items. Each item was characterized either by many or by few features. We identified parameters associated or not associated with menstrual cycle phases. Irrespective of the menstrual cycle phase, women (1) responded faster to living items as well as to nouns characterized by many features compared to non-living items and items characterized by few features, (2) showed higher accuracy to non-living items and items having many features, and (3) showed negative correlation between response time (RT) and theta amplitude. RT, accuracy and post-stimulus theta amplitude were not statistically significantly different among early follicular, late follicular or luteal women. In early follicular but not in late follicular or luteal women, we observed (1) a positive correlation between E2 and latency in RT, (2) a negative correlation between E2 and accuracy, and (3) a negative correlation between E2 and post-stimulus theta amplitude. A mosaic of menstrual cycle phase-dependent and -independent associations may indicate that a similar performance in each menstrual cycle phase is related to different modulation of synaptic activity by hormones.

Consolidation of temporal order in episodic memories.

Even though it is known that sleep benefits declarative memory consolidation, the role of sleep in the storage of temporal sequences has rarely been examined. Thus we explored the influence of sleep on temporal order in an episodic memory task followed by sleep or sleep deprivation. Thirty-four healthy subjects (17 men) aged between 19 and 28 years participated in the randomized, counterbalanced, between-subject design. Parameters of interests were NREM/REM cycles, spindle activity and spindle-related EEG power spectra. Participants of both groups (sleep group/sleep deprivation group) performed retrieval in the evening, morning and three days after the learning night. Results revealed that performance in temporal order memory significantly deteriorated over three days only in sleep deprived participants. Furthermore our data showed a positive relationship between the ratios of the (i) first NREM/REM cycle with more REM being associated with delayed temporal order recall. Most interestingly, data additionally indicated that (ii) memory enhancers in the sleep group show more fast spindle related alpha power at frontal electrode sites possibly indicating access to a yet to be consolidated memory trace. We suggest that distinct sleep mechanisms subserve different aspects of episodic memory and are jointly involved in sleep-dependent memory consolidation.

Pre-stimulus alpha phase-alignment predicts P1-amplitude.

Since years there is a hotly discussed dispute whether event-related potentials are either generated by an evoked component or by resetting of ongoing phase. We argue that phase-reset must not be proven in order to accept the general involvement of phase in ERP-generation as it is only one of several possible mechanisms influencing or generating certain ERP-components. Supporting data are presented showing that positive peaks of ongoing pre-stimulus alpha activity are not randomly distributed in time across trials. Most importantly, we found that a certain kind of pre-stimulus phase concentration that represents a continuous development of an alpha wave up to the time window where the P1 is generated is associated with an enlarged event-related component. We conclude that ongoing oscillations cannot be considered random background noise (even before stimulus onset) and that there are probably more phase-mechanisms that can contribute to the ERP-generation.

Cognitive processes in disorders of consciousness as revealed by EEG time-frequency analyses.

OBJECTIVE: Although behavioral evaluation of awareness in disorders of consciousness is difficult it remains the clinical standard. We believe that the refinement of EEG and analyses techniques would improve our characterization of those patients. METHODS: We focused on cognitive processing in a sample of 12 control subjects, eight vegetative-state patients, and 13 patients in the minimally consciousness state using EEG. We used an 'active paradigm' which asks subjects to follow instructions, specifically to actively count own or other names as compared to passively listening to them. EEG data was then analyzed using an advanced EEG analysis technique. RESULTS: Results revealed that all groups exhibit a stronger theta-synchronization to their own names when forced to count them. We also observed a delay in theta power in response to targets relative to non-targets when participants were instructed to count their own name. CONCLUSION: Active paradigms are able to induce a different oscillatory activity compared to passive paradigms. Differences between controls and the pathologic groups are prominent in the theta- and alpha-band. SIGNIFICANCE: Time-frequency analyses allow to focus on distinct cognitive processes in patients with disorders of consciousness and thereby contribute to a refined understanding of severely brain-injured patients.

Spontaneous locally restricted EEG alpha activity determines cortical excitability in the motor cortex.

There is growing interest in the functional meaning of rhythmical brain activity. For oscillatory brain activity around 10 Hz in the human electroencephalogram (EEG) it is discussed whether it is associated with the level of cortical excitation. However, it is not clear whether the relation between 10 Hz EEG oscillatory activity and cortical excitability is a global, locally very unspecific phenomenon or whether focal 10 Hz oscillations in the human brain are a highly specific correlate of the cortical excitation level. To determine this open question, multichannel EEG was combined with transcranial magnetic stimulation (TMS) applied to the primary motor cortex in this study. The present data showed that a motor evoked potential was elicited more easily when alpha power immediately preceding the magnetic pulse was low, and vice versa. Interestingly, this effect was only found for very local EEG alpha activity at sites overlying the cortical motor areas to which the TMS pulses were applied. This was verified using source localization in 3D space. These data provide evidence that the magnitude of motor cortical excitability is determined by the amount of topographically specific alpha oscillations in the sensorimotor cortex.

Visual P2 component is related to theta phase-locking.

In this study we investigated the hypothesis whether P2-related differences tested in a visual priming paradigm are associated with theta phase-locking. We recorded the EEG from 31 electrodes and calculated phase-locking index and total power differences for frequencies between 2 and 20 Hz. ERPs (event-related potentials) were analyzed for P1, N1 and P2 components. P2 showed strongest task-related amplitude differences between congruent and incongruent targets. A source analyses was performed for the P2 component using sLoreta that revealed local generators of the P2 in parieto-occipital regions. Phase-locking analyses showed specific effects in the theta range (4-6 Hz) appearing in time windows at around the P2 component. We draw the conclusion that phase-locked theta reflect top-down regulation processes mediating information between memory systems and is in part involved in the modulation of the P2 component.

Are event-related potential components generated by phase resetting of brain oscillations? A critical discussion.

The event-related potential (ERP) is one of the most popular measures in human cognitive neuroscience. During the last few years there has been a debate about the neural fundamentals of ERPs. Two models have been proposed: The evoked model states that additive evoked responses which are completely independent of ongoing background electroencephalogram generate the ERP. On the other hand the phase reset model suggests a resetting of ongoing brain oscillations to be the neural generator of ERPs. Here, evidence for either of the two models is presented and validated, and their possible impact on cognitive neuroscience is discussed. In addition, future prospects on this field of research are presented.

Dissociation of sustained attention from central executive functions: local activity and interregional connectivity in the theta range.

Human brain oscillatory activity was analysed in the electroencephalographic theta frequency range (4-7 Hz) while subjects executed complex sequential finger movements with varying task difficulty and memory load. Local frontal-midline theta activity was associated with the general level of cognitive demand, with the highest amplitudes in the most demanding condition. Using low-resolution electromagnetic tomography analysis (LORETA), this theta activity was localized in the anterior cingulate gyrus including the cingulate motor area. These results suggest that local theta activity in the anterior cingulate gyrus represents correlates of an attentional system that allocate cognitive resources. In addition, interregional connectivity in the theta frequency range was modulated by memory-related executive functions independently of task difficulty. Connectivity analyses revealed a more distributed long-range network including frontal and parietal cortices during execution of novel compared with well-trained finger movement sequences. Thus, these results are compatible with a model in which theta long-range coupling indicates integration of sensory information into executive control components of complex motor behaviour.

The significance of sigma neurofeedback training on sleep spindles and aspects of declarative memory.

The functional significance of sleep spindles for overnight memory consolidation and general learning aptitude as well as the effect of four 10-minute sessions of spindle frequency (11.6-16 Hz, sigma) neurofeedback-training on subsequent sleep spindle activity and overnight performance change was investigated. Before sleep, subjects were trained on a paired-associate word list task after having received either neurofeedback training (NFT) or pseudofeedback training (PFT). Although NFT had no significant impact on subsequent spindle activity and behavioral outcomes, there was a trend for enhanced sigma band-power during NREM (stage 2 to 4) sleep after NFT as compared to PFT. Furthermore, a significant positive correlation between spindle activity during slow wave sleep (in the first night half) and overall memory performance was revealed. The results support the view that the considerable inter-individual variance in sleep spindle activity can at least be partly explained by differences in the ability to acquire new declarative information. We conclude that the short NFT before sleep was not sufficient to efficiently enhance phasic spindle activity and/or to influence memory processing. NFT was, however, successful in increasing sigma power, presumably because sigma NFT effects become more easily evident in actually trained frequency bands than in associated phasic spindle activity.

Sleep spindle-related activity in the human EEG and its relation to general cognitive and learning abilities.

Stage 2 sleep spindles have been previously viewed as useful markers for the development and integrity of the CNS and were more currently linked to 'offline re-processing' of implicit as well as explicit memory traces. Additionally, it had been discussed if spindles might be related to a more general learning or cognitive ability. In the present multicentre study we examined the relationship of automatically detected slow (< 13 Hz) and fast (> 13 Hz) stage 2 sleep spindles with: (i) the Raven's Advanced Progressive Matrices (testing 'general cognitive ability'); as well as (ii) the Wechsler Memory scale-revised (evaluating memory in various subdomains). Forty-eight healthy subjects slept three times (separated by 1 week) for a whole night in a sleep laboratory with complete polysomnographic montage. Whereas the first night only served adaptation and screening purposes, the two remaining nights were preceded either by an implicit mirror-tracing or an explicit word-pair association learning or (corresponding) control task. Robust relationships of slow and fast sleep spindles with both cognitive as well as memory abilities were found irrespectively of whether learning occurred before sleep. Based on the present findings we suggest that besides being involved in shaping neuronal networks after learning, sleep spindles do reflect important aspects of efficient cortical-subcortical connectivity, and are thereby linked to cognitive- and memory-related abilities alike.

Oscillatory EEG correlates of episodic trace decay.

Recent studies suggest that human theta oscillations appear to be functionally associated with memory processes. It is less clear, however, to what type of memory sub-processes theta is related. Using a continuous word recognition task with different repetition lags, we investigate whether theta reflects the strength of an episodic memory trace or general processing demands, such as task difficulty. The results favor the episodic trace decay hypothesis and show that during the access of an episodic trace in a time window of approximately 200-400 ms, theta power decreases with increasing lag (between the first and second presentation of an item). LORETA source localization of this early theta lag effect indicates that parietal regions are involved in episodic trace processing, whereas right frontal regions may guide the process of retrieval. We conclude that episodic encoding can be characterized by two different stages: traces are first processed at parietal sites at approximately 300 ms, then further processing takes place in regions of the medial temporal lobe at approximately 500 ms. Only the first stage is related to theta, whereas the second is reflected by a slow wave with a frequency of approximately 2.5 Hz.

Relevance of EEG alpha and theta oscillations during task switching.

In a task switching design, we investigated the question whether long-range theta coupling primarily reflects top-down control processes. Switch and stay trials did not differ with respect to memory load or global working memory (WM) demands. The results revealed significantly stronger theta coupling (in a range of 4-7 Hz) between prefrontal and posterior regions during switch as compared to stay trials. Power differences, reflecting more local effects, were largest in the upper alpha band (10-13 Hz) and over posterior brain areas, possibly reflecting long-term memory activation. The conclusion of the present study is that long-range coherent oscillatory activity in the theta band reflects top-down activation rather than global WM functions.

A shift of visual spatial attention is selectively associated with human EEG alpha activity.

Event-related potentials and ongoing oscillatory electroencephalogram (EEG) activity were measured while subjects performed a cued visual spatial attention task. They were instructed to shift their attention to either the left or right visual hemifield according to a cue, which could be valid or invalid. Thereafter, a peripheral target had to be evaluated. At posterior parietal brain areas early components of the event-related potential (P1 and N1) were higher when the cue had been valid compared with invalid. An anticipatory attention effect was found in EEG alpha magnitude at parieto-occipital electrode sites. Starting 200 ms before target onset alpha amplitudes were significantly stronger suppressed at sites contralateral to the attended visual hemifield than ipsilateral to it. In addition, phase coupling between prefrontal and posterior parietal electrode sites was calculated. It was found that prefrontal cortex shows stronger phase coupling with posterior sites that are contralateral to the attended hemifield than ipsilateral sites. The results suggest that a shift of attention selectively modulates excitability of the contralateral posterior parietal cortex and that this posterior modulation of alpha activity is controlled by prefrontal regions.

Phase synchronization between theta and upper alpha oscillations in a working memory task.

Motivated by findings that theta and upper alpha oscillations respond selectively to different types of memory demands, we investigated the role of phase synchronization in a memory scanning task. During retention, we found a load dependent increase in upper alpha power at O2 and P4 and a significant upper alpha:theta phase synchronization between right posterior, central and left anterior sites. During retrieval, a load dependent increase in upper alpha phase locking was observed at O2 and an increase in upper alpha:theta phase synchronization between right posterior and left anterior sites. We suggest that theta reflects central executive functions whereas upper alpha may be important for the reactivation of long-term memory codes in short-term memory. The interplay between theta and upper alpha may be reflected by phase synchronization between these frequencies.

Intelligence related upper alpha desynchronization in a semantic memory task.

Recent evidence shows that event-related (upper) alpha desynchronization (ERD) is related to cognitive performance. Several studies observed a positive, some a negative relationship. The latter finding, interpreted in terms of the neural efficiency hypothesis, suggests that good performance is associated with a more 'efficient', smaller extent of cortical activation. Other studies found that ERD increases with semantic processing demands and that this increase is larger for good performers. Studies supporting the neural efficiency hypothesis used tasks that do not specifically require semantic processing. Thus, we assume that the lack of semantic processing demands may at least in part be responsible for the reduced ERD. In the present study we measured ERD during a difficult verbal-semantic task. The findings demonstrate that during semantic processing, more intelligent (as compared to less intelligent) subjects exhibited a significantly larger upper alpha ERD over the left hemisphere. We conclude that more intelligent subjects exhibit a more extensive activation in a semantic processing system and suggest that divergent findings regarding the neural efficiency hypotheses are due to task specific differences in semantic processing demands.

Intelligence related differences in EEG-bandpower.

Several studies on the relationship between event-related desynchronization/synchronization (ERD/ERS) and cognitive performance revealed contradictory results particularly for the alpha band. Studies from our laboratory have shown that good performers show a larger upper alpha ERD (interpreted in terms of larger cortical activation) than bad performers. In contrast, other researchers found evidence for the neural efficiency hypothesis, which states that more intelligent subjects exhibit a smaller extent of cortical activation, which is assumed to be reflected by a smaller upper alpha ERD. Here we address the question whether these divergent results may be due to differences in general task difficulty. Using a modified version of the RAVEN, individually divided into easy and difficult tasks, a group of average and a group of highly intelligent subjects (IQ- and IQ+) have been investigated. While in the theta frequency IQ+ subjects generally exhibited a significantly stronger activation, we found a significant interaction of task difficulty and IQ group in the upper alpha band, indicating both, a weaker activation for the high IQ group during the easy tasks, and a significant increase from easy to difficult tasks for IQ+ only.

Frequency characteristics of evoked and oscillatory electroencephalic activity in a human memory scanning task.

Research on memory scanning tasks indicates that oscillatory activity increases with load during retention, whereas evoked power (EP, comprising the P3) decreases during retrieval. We investigate the question, whether both phenomena are primarily related to theta oscillations. However, we found that during retention alpha oscillations increased with load and that exactly this frequency exhibits increased phase locking - measured by a specially developed phase locking index (PLI) - during retrieval. The decreased P3 amplitude was related to decreased delta EP and PLI. The P3 coincides with the last of three evoked alpha peaks. Thus, alpha may be important for the timing of the scanning and the evaluation of the read out process that most likely is manifested by the P3.

The interplay between theta and alpha oscillations in the human electroencephalogram reflects the transfer of information between memory systems.

The exchange of information between the working and long-term memory system (WMS and LTMS) was investigated. We analyzed evoked theta and upper alpha desynchronization in a special memory task, designed to study the transfer of information between both memory systems. The results show that during attempts to retrieve information from the LTMS, evoked theta oscillations spread from anterior to posterior recording sites. When information actually is retrieved, the direction reverses and theta spreads to frontal sites. This time point--when direction reverses--varies between subjects to a large extent but is significantly correlated with memory performance and the onset of upper alpha desynchronization. We conclude that this phenomenon reflects the transfer of information between the WMS and LTMS