Oscillatory EEG correlates of arithmetic strategy use in addition and subtraction

Adults use different strategies in mental arithmetic. For example, they directly retrieve the answer from memory or calculate by means of procedural strategies. Despite growing insight into the hemodynamic and electrophysiological correlates of these strategies, the functional changes in the oscillatory brain dynamics during the use of these strategies remain unknown. In the present high-resolution electroencephalography (EEG) study, we analysed event-related synchronisation (ERS) and desynchronisation (ERD) in the theta and alpha bands while participants solved addition and subtraction problems, which displayed a high probability of retrieval or procedural strategy use. Findings revealed that arithmetic fact retrieval is reflected in left-hemispheric ERS in the theta band, whereas the application of procedural strategies is accompanied by bilateral parietooccipital ERD in the alpha band. The topographical and frequency specificity of the strategy effects provides a start for the development of electrophysiological indices of strategy use in arithmetic. B. De Smedt and R. H. Grabner equally contributed to this study and should be regarded as joint first authors.     

Differences in induced gamma and upper alpha oscillations in the human brain related to verbal/performance and emotional intelligence.

Participating in the study were 30 respondents, who could be clustered as high-average verbal/performance intelligent (HIQ/AIQ), or emotionally intelligent (HEIQ/AEIQ). The EEG was recorded while students were performing two tasks: the Raven's advanced progressive matrices (RAPM), and identifying emotions in pictures (IDEM). Significant differences in event-related desynchronization/synchronization (ERD/ERS) related to verbal/performance intelligence were only observed while respondents solved the RAPM. The HIQ and AIQ groups displayed temporal and spatial differently induced gamma band activity. Significant differences in ERD/ERS related to emotional intelligence were only observed for the IDEM task. HEIQ individuals displayed more gamma band ERS and less upper alpha band ERD than did AEIQ individuals. It can be concluded that HIQ and HEIQ individuals employed more adequate strategies for solving the problems at hand. The results further suggest that emotional intelligence and verbal/performance intelligence represent distinct components of the cognitive architecture.     

Electrophysiological correlates of memory processing in early Finnish–Swedish bilinguals

Event-related desynchronization (ERD) and event-related synchronization (ERS) of the 1–30 EEG frequencies were studied in eight early Finnish–Swedish bilinguals during an auditory bilingual Sternberg memory task using Finnish–Swedish cognates as stimuli. Only subtle differences between languages were expected, since cognates have been assumed to have shared conceptual representations in the bilingual memory. Encoding elicited theta and alpha frequency ERS and beta frequency ERD responses in both languages. Retrieval elicited theta ERS and alpha and beta ERD responses. Some statistically significant differences between encoding and retrieval in Finnish versus Swedish emerged: greater theta and alpha ERS responses were observed during encoding in Swedish than during encoding in Finnish. During between-language retrieval, later-appearing theta ERS and alpha ERD responses were elicited as compared to within-language retrieval. These delayed oscillatory responses might reflect the involvement of central executive attentional functions in relation to language switching.     

Effects of normal aging on event-related desynchronization/synchronization during a memory task in humans

Event-related desynchronization (ERD) and event-related synchronization (ERS) of the 1-20Hz EEG frequencies were studied using wavelet transforms in young (n = 10, mean age 22) and elderly subjects (n = 10, mean age 65) performing an auditory Sternberg memory task with words as stimuli. In both age groups, encoding of the four-word memory set elicited ERS in the theta and alpha frequency range. Theta ERS, and ERD in the alpha and beta bands were observed during retrieval. During encoding, the elderly showed greater alpha ERS and smaller theta ERS. During retrieval, smaller alpha ERD and theta ERS was found in the elderly subjects. Also, in the elderly, beta ERD was elicited in the late time window during retrieval. The statistically significant differences between the age groups were more marked during retrieval than during encoding. The results indicate that although the two groups performed equally well behaviorally in the task and the elderly subjects were cognitively intact, normal aging affects oscillatory theta, alpha and beta responses particularly during retrieval from working memory. The ERD/ERS patterns of the elderly resemble those of children found in a recent study, which might suggest that those memory-related brain processes that evolve later in childhood are the first to be affected in older age.     

Brain oscillatory 1-30 Hz EEG ERD/ERS responses during the different stages of an auditory memory search task

Event-related desynchronization (ERD) and event-related synchronization (ERS) responses of 1-30 Hz EEG frequencies during the different stages of an auditory Sternberg memory task were examined. The ERD/ERS responses were examined separately for successive memory set items (four) and for the two recognition conditions (YES/NO). The presentation of the memory set elicited ERS responses in the theta and alpha frequencies, and also beta ERD responses. These ERD/ERS responses elicited during encoding were found to evolve with successive memory set item presentation. The ERD/ERS responses elicited during the presentation of the probe dissociated significantly between the two recognition conditions (YES/NO). When the probe was included in the memory set (YES condition), recognition elicited stronger alpha and beta frequency ERD responses as compared to the NO condition. The findings from the current study verify that alpha ERD/ERS responses robustly dissociate between auditory encoding and recognition. The increasing alpha ERS responses with increasing memory set item presentation during encoding may be correlates of the functioning phonological loop, active memory maintenance and/or attention. The alpha ERD responses during recognition are undoubtedly associated with auditory memory search processes and distinguish between previously presented versus not presented verbal material. We propose that alpha ERD/ERS responses reflect explicitly auditory memory processes, discriminating between auditory encoding and recognition. Theta ERS responses may be associated with working memory processes, and possibly more specifically with the functioning of the central executive. Beta ERD/ERS responses may reflect also cognitive and/or memory processing, rather than merely the activity of the motor cortices.     

Brain oscillation and connectivity during a chemistry visual working memory task

Many studies have reported that frontal theta and posterior alpha activities are associated with working memory tasks. However, fewer studies have focused on examining whether or not the frontal alpha or posterior theta can play a role in the working memory task. This study investigates electroencephalography (EEG) dynamics and connectivity among different brain regions' theta and alpha oscillations. The EEG was collected from undergraduate students (n = 64) while they were performing a Sternberg-like working memory task involving chemistry concepts. The results showed that the frontal midline cluster exhibited sustained theta augmentation across the periods of stimulus presentations, maintenance, and probe presentation, suggesting that the frontal midline theta might associate with facilitating the central execute function to maintain information in the working memory. Study of the central parietal and the occipital clusters revealed a sequence of theta augmentation followed by alpha suppression at constant intervals after the onset of stimulus and probe presentations, suggesting that the posterior theta might be associated with sensory processing, theta gating, or stimulus selection. It further suggests that the posterior alpha event-related de-synchronization (ERD) might be linked to direct information flow into and out of the long-term memory (LTM) and precede stimulus recognition. An alternating phasic alpha event-related synchronization (ERS) and ERD following the 1st stimulus and probe presentations were observed at the occipital cluster, in which alpha ERS might be linked to the inhibition of irrelevant information.     

Oscillatory EEG Changes During Arithmetic Learning in Children

Most studies have investigated brain activation changes after the course of arithmetic learning, and the question remains whether these changes are detectable during the course of learning, i.e., before memory consolidation. Twenty-four fifth graders solved multiplication problems while ongoing electroencephalography (EEG) was recorded. The arithmetic training revealed reduced errors together with a power increase in theta (4–7 Hz) but not in lower alpha (8–10 Hz) or upper alpha (10–13 Hz) bands. We conclude that increases in theta power subserved a shift from slow, procedural strategies to more efficient, automated procedural and retrieval strategies, which led to more efficient performance.     

Event‐related EEG time‐frequency PCA and the orienting reflex to auditory stimuli

We recently reported an auditory habituation series with counterbalanced indifferent and significant (counting) instructions. Time‐frequency (t‐f) analysis of electrooculogram‐corrected EEG was used to explore event‐related synchronization (ERS)/desynchronization (ERD) in four EEG bands using arbitrarily selected time epochs and traditional frequency ranges. ERS in delta, theta, and alpha, and subsequent ERD in theta, alpha, and beta, showed substantial decrement over trials, yet effects of stimulus significance (count vs. no‐task) were minimal. Here, we used principal components analysis (PCA) of the t‐f data to investigate the natural frequency and time combinations involved in such stimulus processing. We identified four ERS and four ERD t‐f components: six showed decrement over trials, four showed count > no‐task effects, and six showed Significance × Trial interactions. This increased sensitivity argues for the wider use of our data‐driven t‐f PCA approach.     

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.     

The Functional Significance of Absolute Power with Respect to Event-Related Desynchronization

The question is examined whether the extent of changes in relative band power as measured by event-related desynchronization (ERD) depends on absolute band power. The results for target stimuli of a simple oddball task indicate that the prestimulus (reference) level of absolute band power has indeed a strong influence on ERD. Whereas for the alpha band large band power in the reference interval is related to a strong degree of alpha suppression as measured by ERD, the opposite holds true for the theta band. Here, a low level of band power during the reference interval is related to a pronounced increase in band power during the processing of the target stimulus. In contrast to alpha and theta, ERD in the delta band is not influenced by the magnitude of band power in the reference interval.     

Event-related desynchronization/synchronization in the putamen. An SEEG case study

Event-related desynchronization (ERD) and synchronization (ERS) were studied during the invasive exploration of an epileptic surgery candidate. An electrode that was targeted in the amygdalo-hippocampal complex passed through the putamen with several contacts. During a simple self-paced motor task, we observed in the putamen a power decline (ERD) in both the alpha and beta frequency bands, and a rebound phenomenon (ERS) in the beta frequency band, concurrent with the movement of each hand. This is the first report of ERD/ERS in the basal ganglia. Electronic Publication     

Spectra-temporal patterns underlying mental addition: An ERP and ERD/ERS study

Functional neuroimaging data have shown that mental calculation involves fronto-parietal areas that are composed of different subsystems shared with other cognitive functions such as working memory and language. Event-related potential (ERP) analysis has also indicated sequential information changes during the calculation process. However, little is known about the dynamic properties of oscillatory networks in this process. In the present study, we applied both ERP and event-related (de-)synchronization (ERS/ERD) analyses to EEG data recorded from normal human subjects performing tasks for sequential visual/auditory mental addition. Results in the study indicate that the late positive components (LPCs) can be decomposed into two separate parts. The earlier element LPC1 (around 360 ms) reflects the computing attribute and is more prominent in calculation tasks. The later element LPC2 (around 590 ms) indicates an effect of number size and appears larger only in a more complex 2-digit addition task. The theta ERS and alpha ERD show modality-independent frontal and parietal differential patterns between the mental addition and control groups, and discrepancies are noted in the beta ERD between the 2-digit and 1-digit mental addition groups. The 2-digit addition (both visual and auditory) results in similar beta ERD patterns to the auditory control, which may indicate a reliance on auditory-related resources in mental arithmetic, especially with increasing task difficulty. These results coincide with the theory of simple calculation relying on the visuospatial process and complex calculation depending on the phonological process.     

Computational model of thalamo-cortical networks: dynamical control of alpha rhythms in relation to focal attention.

EEG/MEG rhythmic activities such as alpha rhythms, of the visual or of the somato-sensory cortex, are commonly modulated as subjects perform certain tasks or react to specific stimuli. In general, these activities change depending on extrinsic or intrinsic events. A decrease of the amplitude of alpha rhythmic activity occurring after a given event, which manifests as a decrease of a spectral peak, is called event-related desynchronization (ERD), whereas the inverse is called event-related synchronization (ERS), since it is assumed that the power of a spectral peak is related to the degree of synchrony of the underlying oscillating neuronal populations. An intriguing observation in this respect [Pfurtscheller and Neuper, Neurosci. Lett. 174 (1994) 93-96] was that ERD of alpha rhythms recorded over the central areas was accompanied by ERS, within the same frequency band, recorded over neighboring areas. In case the event was a hand movement, ERD was recorded over the scalp overlying the hand cortical area, whereas ERS was concomitantly recorded over the midline, whereas if the movement was of the foot the opposite was found. We called this phenomenon 'focal ERD/surround ERS'. The question of how this phenomenon may be generated was approached by means of a computational model of thalamo-cortical networks, that incorporates basic properties of neurons and synaptic interactions. These simulation studies revealed that this antagonistic ERD/ERS phenomenon depends on the functional interaction between the populations of thalamo-cortical cells (TCR) and reticular nucleus cells (RE) and on how this interaction is modulated by cholinergic inputs. An essential feature of this interaction is the existence of cross-talk between different sectors of RE that correspond to distinct sensory modules (e.g. hand, foot). These observations led us to formulate the hypothesis that this basic neurophysiological mechanism can account for the general observation that enhanced attention given to a certain stimulus (the focus) is coupled to inhibition of attention to other stimuli (the surround).     

Electrophysiological markers of skill-related neuroplasticity

Neuroplasticity involved in acquiring a new cognitive skill was investigated with standard time domain event-related potentials (ERPs) of scalp-recorded electroencephalographic (EEG) activity and frequency domain analysis of EEG oscillations looking at the event-related synchronization (ERS) and desynchronization (ERD) of neural activity. Electroencephalographic activity was recorded before and after practice, while participants performed alphabet addition (i.e., E + 3 = G, true or false?). Participant's performance became automated with practice through a switch in cognitive strategy from mentally counting-up in the alphabet to retrieving the answer from memory. Time domain analysis of the ERPs revealed a prominent positive peak at ∼300 ms that was not reactive to problem attributes but was reduced with practice. A second prominent positive peak observed at ∼500 ms was found to be larger after practice, mainly for problems presented with correct answers. Frequency domain spectral analyses yielded two distinct findings: (1) a frontal midline ERS of theta activity that was greater after practice, and (2) a beta band ERD that increased with problem difficulty before, but not after practice. Because the EEG oscillations were not phase locked to the stimulus, they were viewed as being independent of the time domain results. Consequently, use of time and frequency domain analyses provides a more comprehensive account of the underlying electrophysiological data than either method alone. When used in combination with a well-defined cognitive/behavioral paradigm, this approach serves to constrain the interpretations of EEG data and sets a new standard for studying the neuroplasticity involved in skill acquisition.     

Event-related dynamics of cortical rhythms: frequency-specific features and functional correlates.

Oscillations in the alpha and beta band (<35 Hz) display a dynamic behavior and show characteristic spatiotemporal patterns in sensory, motor and cognitive tasks. The event-related desynchronization (ERD) of alpha band and beta rhythms can be seen as a correlate of an activated cortical area with an increased excitability level of neurons. An event-related synchronization (ERS) of frequency components between 10 and 13 Hz may represent a deactivated cortical area or inhibited cortical network, at least under certain circumstances. It is hypothesized, that antagonistic ERD/ERS patterns, called 'focal ERD/surround ERS', may reflect a thalamo-cortical mechanism to enhance focal cortical activation by simultaneous inhibition of other cortical areas. Induced oscillations in the beta band (13-35 Hz, beta ERS) were found in sensorimotor areas after voluntary movement and after somatosensory stimulation. This may be interpreted as a state of 'inhibition' of neural circuitry in the primary motor cortex. Simultaneous activation of the motor cortex by e.g. motor imagery lead to an attenuation of the beta ERS. Moreover, there is evidence that the frequency of the induced beta oscillations represent a 'resonance-like frequency' of underlying cortical networks. However, further research is needed to investigate the functional meaning of bursts of beta oscillations below 35 Hz.     

Variability and Reliability of Paired-Pulse Depression and Cortical Oscillation Induced by Median Nerve Stimulation

Paired-pulse depression (PPD) has been widely used to investigate the functional profiles of somatosensory cortical inhibition. However, PPD induced by somatosensory stimulation is variable, and the reasons for between- and within-subject PPD variability remains unclear. Therefore, the purpose of this study was to clarify the factors influencing PPD variability induced by somatosensory stimulation. The study participants were 19 healthy volunteers. First, we investigated the relationship between the PPD ratio of each component (N20m, P35m, and P60m) of the somatosensory magnetic field, and the alpha, beta, and gamma band changes in power [event-related desynchronization (ERD) and event-related synchronization (ERS)] induced by median nerve stimulation. Second, because brain-derived neurotrophic factor (BDNF) gene polymorphisms reportedly influence the PPD ratio, we assessed whether BDNF genotype influences PPD ratio variability. Finally, we evaluated the test–retest reliability of PPD and the alpha, beta, and gamma ERD/ERS induced by somatosensory stimulation. Significant positive correlations were observed between the P60m_PPD ratio and beta power change, and the P60m_PPD ratio was significantly smaller for the beta ERD group than for the beta ERS group. P35m_PPD was found to be robust and highly reproducible; however, P60m_PPD reproducibility was poor. In addition, the ICC values for alpha, beta, and gamma ERD/ERS were 0.680, 0.760, and 0.552 respectively. These results suggest that the variability of PPD for the P60m deflection may be influenced by the ERD/ERS magnitude, which is induced by median nerve stimulation.     

Auditory processing indexed by stimulus-induced alpha desynchronization in children.

By means of magnetoencephalography (MEG), we investigated event-related synchronization and desynchronization (ERS/ERD) in auditory cortex activity, recorded from twelve children aged four to six years, while they passively listened to a violin tone and a noise-burst stimulus. Time-frequency analysis using Wavelet Transform was applied to single-trials of source waveforms observed from left and right auditory cortices. Stimulus-induced changes in non-phase-locked activities were evident. ERS in the beta range (13-30 Hz) lasted only for 100 ms after stimulus onset. This was followed by prominent alpha ERD, which showed a clear dissociation between the upper (12 Hz) and lower (8 Hz) alpha range in both left and right auditory cortices for both stimuli. The time courses of the alpha ERD (onset around 300 ms, peak at 500 ms, offset after 1500 ms) were similar to those previously found for older children and adults with auditory memory related tasks. For the violin tone only, the ERD lasted longer in the upper than the lower alpha band. The findings suggest that induced alpha ERD indexes auditory stimulus processing in children without specific cognitive task requirement. The left auditory cortex showed a larger and longer-lasting upper alpha ERD than did the right auditory cortex, likely reflecting hemispheric differences in maturational stages of neural oscillatory mechanisms.     

A frequency band analysis of two-year-olds' memory processes

Research on the functional meaning of EEG frequency bands during memory processing has only examined two developmental periods: infancy and from late childhood to adulthood. The purpose of this study was to examine changes in EEG power for three toddler EEG frequency bands (3-5Hz, 6-9Hz, 10-12Hz) during a verbal recall task. To this end, we asked three questions: (a) Which frequency band(s) discriminate baseline from memory processing?; (b) Which frequency band(s) differentiate between memory encoding and retrieval processes?; (c) Which frequency band(s) distinguish toddlers with high and low verbal recall performance? Analysis of 2-year-olds' (n=79) power values revealed that all three frequency bands differentiated the retrieval and encoding phases from the baseline phase; however, the particular regions that exhibited this dissociation varied. Retrieval-related increases in 3-5Hz (theta) power were widespread. Only the 3-5Hz and 6-9Hz bands distinguished encoding and retrieval processes; retrieval power values were higher than encoding power values. High and low verbal recall performers were discriminated by all frequency bands; high performers had greater power values than low performers. Thus, the 3-5Hz (theta) and 6-9Hz (alpha) bands were most informative about 2-year-olds' memory processes. Theta and alpha rhythms are critical to memory processes during late childhood and adulthood, and our findings provide initial evidence that these rhythms are also intricately linked to memory processing during toddlerhood. These findings are discussed in relation to behavioral changes in memory processes.     

Cortical processing of vowels and tones as measured by event-related desynchronization

Summary Event-Related Desynchronization (ERD) and Synchronization (ERS) were studied in 20 normal subjects during a Sternberg-type auditory memory-scanning paradigm. Half of the subjects performed the experiment with vowels and the other half with tones as stimuli. The stimuli consisted of 100 msec long synthesized vowels and 100 msec long tones produced by eight different synthesized instruments. In this paradigm each trial started with the presentation of a visual warning signal, after which a four-stimulus set was presented for memorization whereafter a probe stimulus was presented and identified by the subject as belonging or not belonging to the memorized set. The ERD/ERS of the lower (8–10 Hz) and upper (10–12 Hz) alpha frequency bands differed in their reactivity to stimulus type; the differences between the two frequency bands reached statistical significance only in the case of vowels. The presentation of the memory set elicited ERS which was more pronounced in the 10–12 Hz frequency band and greater for vowels than for tones. On the other hand, the presentation of the probe elicited ERD which was greater for vowels than for tones, especially in the upper alpha frequency band. The results of this exploratory study suggest that ERD is closely related to memory processes and that the ERD/ERS-technique might provide a valuable tool for future reseach encompassing more complex auditory stimulation like speech and music.This study was financially supported by the Council for Social Sciences, Academy of Finland (project 7338).     

The Influence of Mozart’s Sonata K. 448 on Brain Activity During the Performance of Spatial Rotation and Numerical Tasks

Summary: The study investigated the influence of Mozart’s music on respondents’ brain activity while solving spatial rotation and numerical tasks. The method of induced event-related desynchronization/synchronization (ERD/ERS) and coherence (ERCoh) was used. The music condition had a beneficial influence on respondents’ performance of spatial rotation tasks, and a slightly negative influence on the performance of numerical tasks as compared with the silence condition. On the psychophysiological level a general effect of Mozart’s music on brain activity in the induced gamma band was observed, accompanied by a more specific effect in the induced lower-2 alpha band which was only present while respondents solved the numerical tasks. It is suggested that listening to Mozart’s music increases the activity of specific brain areas and in that way facilitates the selection and “binding” together of pertinent aspects of sensory stimulus into a perceived whole.