Alpha Suppression Research Articles

Cognitive training, but not EEG-neurofeedback, improves working memory in healthy volunteers.

Working memory performance can be influenced by motivational factors, which may be associated with specific brain activities, including suppression of alpha oscillations. We investigated whether providing individuals online feedback about their ongoing oscillations (EEG-neurofeedback) can improve working memory under high and low reward expectancies. We combined working memory training with neurofeedback to enhance alpha suppression in a monetary-rewarded delayed match-to-sample task for visual objects. Along with alpha, we considered the neighbouring theta and beta bands. In a double-blind experiment, individuals were trained over 5 days to suppress alpha power by receiving real-time neurofeedback or control neurofeedback (placebo) in reward and no-reward trials. We investigated (i) whether neurofeedback enhances alpha suppression, (ii) whether monetary reward enhances alpha suppression and working memory, and (iii) whether any performance benefits of neurofeedback-training would transfer to unrelated cognitive tasks. With the same experimental design, we conducted two studies with differing instructions given at the maintenance, yielding together 300 EEG recording sessions. In Study I, participants were engaged in a mental calculation task during maintenance. In Study II, they were instructed to visually rehearse the sample image. Results from Study I demonstrated a significant training and reward-anticipation effect on working memory accuracy and reaction times over 5 days. Neurofeedback and reward anticipation showed effects on theta suppression but not on alpha suppression. Moreover, a cognitive training effect was observed on beta suppression. Thus, neurofeedback-training of alpha was unrelated to working memory performance. Study II replicated the training and reward-anticipation effect on working memory but without any effects of neurofeedback-training on oscillations or working memory. Neither study showed transfer effects of either working memory or neurofeedback-training. A linear mixed-effect model analysis of neurofeedback-independent training-related improvement of working memory combining both studies showed that improved working memory performance was related to oscillatory changes over training days in the encoding and maintenance phases. Improvements in accuracy were related to increasing beta amplitude in reward trials over right parietal electrodes. Improvements in reaction times were related to increases in right parietal theta amplitude during encoding and increased right parietal and decreased left parietal beta amplitudes during maintenance. Thus, while our study provided no evidence that neurofeedback targeting alpha improved the efficacy of working memory training or evidence for transfer, it showed a relationship between training-related changes in parietal beta oscillations during encoding and improvements in accuracy. Right parietal beta oscillations could be an intervention target for improving working memory accuracy.

Dynamics of alpha suppression index both modality specific and general attention processes

EEG alpha power varies under many circumstances requiring visual attention. However, mounting evidence indicates that alpha may not only serve visual processing, but also the processing of stimuli presented in other sensory modalities, including hearing. We previously showed that alpha dynamics during an auditory task vary as a function of competition from the visual modality (Clements et al., 2022) suggesting that alpha may be engaged in multimodal processing. Here we assessed the impact of allocating attention to the visual or auditory modality on alpha dynamics at parietal and occipital electrodes, during the preparatory period of a cued-conflict task. In this task, bimodal precues indicated the modality (vision, hearing) relevant to a subsequent reaction stimulus, allowing us to assess alpha during modality-specific preparation and while switching between modalities. Alpha suppression following the precue occurred in all conditions, indicating that it may reflect general preparatory mechanisms. However, we observed a switch effect when preparing to attend to the auditory modality, in which greater alpha suppression was elicited when switching to the auditory modality compared to repeating. No switch effect was evident when preparing to attend to visual information (although robust suppression did occur in both conditions). In addition, waning alpha suppression preceded error trials, irrespective of sensory modality. These findings indicate that alpha can be used to monitor the level of preparatory attention to process both visual and auditory information, and support the emerging view that alpha band activity may index a general attention control mechanism used across modalities.

Human Brain Dynamics and Coordination Reflect the Task Difficulty of Optical Image Relational Reasoning.

Despite advances in neuroscience, the mechanisms by which human brain resolve optical image formation through relational reasoning remain unclear, particularly its relationships with task difficulty. Therefore, this study explores the underlying brain dynamics involved in optical image formation tasks at various difficulty levels, including those with a single convex lens and a single mirror. Compared to single convex lens relational reasoning with high task difficulty, the single mirror relational reasoning exhibited significantly higher response accuracy and shorter latency. As compared to single mirror tasks, single convex tasks exhibited greater frontal midline theta augmentation and right parietal alpha suppression during phase I and earlier phase II, and augmentation of frontal midline theta, right parietal-occipital alpha, and left mu alpha suppression during late phase II. Moreover, the frontal midline theta power in late phase II predicts the likelihood of solving single convex tasks the best, while the parietal alpha power in phase I is most predictive. In addition, frontal midline theta power exhibited stronger synchronization with right parietal alpha, right occipital alpha, and mu alpha power when solving single convex tasks than single mirror tasks. In summary, having stronger brain dynamics and coordination is vital for achieving optical image formation with greater difficulty.

Alpha Suppression Is Associated with the Tip-of-the-Tongue (TOT) State Whereas Alpha Expression Is Associated with Knowing That One Does Not Know.

The tip-of-the-tongue (TOT) state is a spontaneously occurring metacognitive state that indicates that the answer to a query is almost, but not quite, at hand, i.e., that resolution is imminent. Since the time of William James, a distinctive feeling of nagging frustration has been observed to be associated with TOT states. On a more positive note, TOT states are also associated with intense goal-directed curiosity and with a strong desire to know that translates into successful mental action. The present study showed that prior to the presentation of resolving feedback to verbal queries-if the individual was in a TOT state-alpha suppression was in evidence in the EEG. This alpha suppression appears to be a marker of a spontaneously occurring, conscious, and highly motivating goal-directed internal metacognitive state. At the same time, alpha expression in the same time period was associated with the feeling of not knowing, indicating a more discursive state. Both alpha and alpha suppression were observed broadly across centro-parietal scalp electrodes and disappeared immediately upon presentation of the resolving feedback. Analyses indicated that the occurrence of alpha suppression was associated with participants' verbal affirmations of being in a TOT state, which is also related to subsequent expression of a late positivity when feedback is provided, and to enhanced memory.

Tactile expectancy modulates occipital alpha oscillations in early blindness

Alpha oscillatory activity is thought to contribute to visual expectancy through the engagement of task-relevant occipital regions. In early blindness, occipital alpha oscillations are systematically reduced, suggesting that occipital alpha depends on visual experience. However, it remains possible that alpha activity could serve expectancy in non-visual modalities in blind people, especially considering that previous research has shown the recruitment of the occipital cortex for non-visual processing. To test this idea, we used electroencephalography to examine whether alpha oscillations reflected a differential recruitment of task-relevant regions between expected and unexpected conditions in two haptic tasks (texture and shape discrimination). As expected, sensor-level analyses showed that alpha suppression in parieto-occipital sites was significantly reduced in early blind individuals compared with sighted participants. The source reconstruction analysis revealed that group differences originated in the middle occipital cortex. In that region, expected trials evoked higher alpha desynchronization than unexpected trials in the early blind group only. Our results support the role of alpha rhythms in the recruitment of occipital areas in early blind participants, and for the first time we show that although posterior alpha activity is reduced in blindness, it remains sensitive to expectancy factors. Our findings therefore suggest that occipital alpha activity is involved in tactile expectancy in blind individuals, serving a similar function to visual anticipation in sighted populations but switched to the tactile modality. Altogether, our results indicate that expectancy-dependent modulation of alpha oscillatory activity does not depend on visual experience. Significance statementAre posterior alpha oscillations and their role in expectancy and anticipation dependent on visual experience? Our results show that tactile expectancy can modulate posterior alpha activity in blind (but not sighted) individuals through the engagement of occipital regions, suggesting that in early blindness, alpha oscillations maintain their proposed role in visual anticipation but subserve tactile processing. Our findings bring a new understanding of the role that alpha oscillatory activity plays in blindness, contrasting with the view that alpha activity is task unspecific in blind populations.

The new wireless EEG device Mentalab Explore is a valid and reliable system for the measurement of resting state EEG spectral features

Technological advancements in neuroscience have provided many conveniences to scientists, researchers, and consumers. The emerging electroencephalography (EEG) devices are good examples for it. The objective of this study was to investigate the validity and reliability of a new wireless EEG device Mentalab Explore for resting-state EEG (rsEEG) recordings in eyes open and eyes closed conditions. Twenty-three healthy subjects were recruited for the study. The subjects visited the laboratory on two occasions. On the first day, both devices were used to record rsEEG data, and after 24 h, only the Mentalab Explore was used to record rsEEG for test–retest reliability analysis. We compared the alpha peak frequency, suppression, and mean power between the two devices. Intraclass correlation coefficient was calculated for test–retest reliability analysis. Power spectral density (PSD) was calculated using Welch method. The PSD (eyes closed p < 0.0001; eyes open p = 0.01–0.0001), alpha peak frequency (p < 0.0001), and alpha suppression (p = 0.002–0.0001) from the two EEG devices as well as the test–retest results from the Mentalab device were significantly correlated. There were no significant differences in alpha peak frequency or suppression between the clinical-grade and the new devices for O1 and O2 channels in eyes-closed condition. The two measurements completed ∼24 h apart using the Mentalab system were similar for all the variables. We showed that the consumer-grade Mentalab Explore with gel electrodes is a reliable and valid EEG recording device for rsEEG spectral features.

Neurodynamic correlates for the cross-frequency coupled transcranial alternating current stimulation during working memory performance.

Transcranial current stimulation is a neuromodulation technique used to modulate brain oscillations and, in turn, to enhance human cognitive function in a non-invasive manner. This study investigated whether cross-frequency coupled transcranial alternating current stimulation (CFC-tACS) improved working memory performance. Participants in both the tACS-treated and sham groups were instructed to perform a modified Sternberg task, where a combination of letters and digits was presented. Theta-phase/high-gamma-amplitude CFC-tACS was administered over electrode F3 and its four surrounding return electrodes (Fp1, Fz, F7, and C3) for 20 min. To identify neurophysiological correlates for the tACS-mediated enhancement of working memory performance, we analyzed EEG alpha and theta power, cross-frequency coupling, functional connectivity, and nodal efficiency during the retention period of the working memory task. We observed significantly reduced reaction times in the tACS-treated group, with suppressed treatment-mediated differences in frontal alpha power and unidirectional Fz-delta-phase to Oz-high-gamma-amplitude modulation during the second half of the retention period when network analyses revealed tACS-mediated fronto-occipital dissociative neurodynamics between alpha suppression and delta/theta enhancement. These findings indicate that tACS modulated top-down control and functional connectivity across the fronto-occipital regions, resulting in improved working memory performance. Our observations are indicative of the feasibility of enhancing cognitive performance by the CFC-formed tACS.

Type of bilingualism conditions individual differences in the oscillatory dynamics of inhibitory control.

The present study uses EEG time-frequency representations (TFRs) with a Flanker task to investigate if and how individual differences in bilingual language experience modulate neurocognitive outcomes (oscillatory dynamics) in two bilingual group types: late bilinguals (L2 learners) and early bilinguals (heritage speakers—HSs). TFRs were computed for both incongruent and congruent trials. The difference between the two (Flanker effect vis-à-vis cognitive interference) was then (1) compared between the HSs and the L2 learners, (2) modeled as a function of individual differences with bilingual experience within each group separately and (3) probed for its potential (a)symmetry between brain and behavioral data. We found no differences at the behavioral and neural levels for the between-groups comparisons. However, oscillatory dynamics (mainly theta increase and alpha suppression) of inhibition and cognitive control were found to be modulated by individual differences in bilingual language experience, albeit distinctly within each bilingual group. While the results indicate adaptations toward differential brain recruitment in line with bilingual language experience variation overall, this does not manifest uniformly. Rather, earlier versus later onset to bilingualism—the bilingual type—seems to constitute an independent qualifier to how individual differences play out.

Load-dependent alpha suppression is related to working memory capacity for numbers

Alpha suppression is proposed to reflect a surge in cortical excitability to enhance stimulus processing in working memory. The attenuated state of alpha might reflect the prioritisation of behaviourally relevant information, making it a proxy for working memory functioning. Despite the growing interest in utilising the advancement of brain-based measures to evaluate individuals’ cognitive processes, there was a lack of consistent evidence on the relationship between alpha suppression and working memory performance. To investigate whether interindividual differences in alpha suppression might be related to variability in working memory capacity, we recorded participants’ electroencephalography (EEG) while they performed an arithmetic task of either low or high working memory load. Participants were required to calculate either the product of digits (i.e., low-load condition) or the difference between the product of digits (i.e., high-load condition). We found alpha suppression at parietal regions, which became more prominent as working memory load increased. The pattern was present in approximately 80% of the participants. Importantly, the more the alpha suppressed as working memory load increased, the larger the drops in behavioural performance and the lower the Digit Span score. That is, alpha suppression was more prominent in participants of poor working memory capacity. Our findings suggest that alpha activity, subject to interindividual differences in sensitivity, could serve as a brain-based measure of an individual’s working memory functioning.

Gamma Activation and Alpha Suppression within Human Auditory Cortex during a Speech Classification Task.

The dynamics of information flow within the auditory cortical hierarchy associated with speech processing and the emergence of hemispheric specialization remain incompletely understood. To study these questions with high spatiotemporal resolution, intracranial recordings in 29 human neurosurgical patients of both sexes were obtained while subjects performed a semantic classification task. Neural activity was recorded from posteromedial portion of Heschl's gyrus (HGPM) and anterolateral portion of Heschl's gyrus (HGAL), planum temporale (PT), planum polare, insula, and superior temporal gyrus (STG). Responses to monosyllabic words exhibited early gamma power increases and a later suppression of alpha power, envisioned to represent feedforward activity and decreased feedback signaling, respectively. Gamma activation and alpha suppression had distinct magnitude and latency profiles. HGPM and PT had the strongest gamma responses with shortest onset latencies, indicating that they are the earliest auditory cortical processing stages. The origin of attenuated top-down influences in auditory cortex, as indexed by alpha suppression, was in STG and HGAL. Gamma responses and alpha suppression were typically larger to nontarget words than tones. Alpha suppression was uniformly greater to target versus nontarget stimuli. Hemispheric bias for words versus tones and for target versus nontarget words, when present, was left lateralized. Better task performance was associated with increased gamma activity in the left PT and greater alpha suppression in HGPM and HGAL bilaterally. The prominence of alpha suppression during semantic classification and its accessibility for noninvasive electrophysiologic studies suggests that this measure is a promising index of auditory cortical speech processing.SIGNIFICANCE STATEMENT Understanding the dynamics of cortical speech processing requires the use of active tasks. This is the first comprehensive intracranial electroencephalography study to examine cortical activity within the superior temporal plane, lateral superior temporal gyrus, and the insula during a semantic classification task. Distinct gamma activation and alpha suppression profiles clarify the functional organization of feedforward and feedback processing within the auditory cortical hierarchy. Asymmetries in cortical speech processing emerge at early processing stages. Relationships between cortical activity and task performance are interpreted in the context of current models of speech processing. Results lay the groundwork for iEEG studies using connectivity measures of the bidirectional information flow within the auditory processing hierarchy.

Attenuated alpha oscillation and hyperresponsiveness reveals impaired perceptual learning in migraineurs

BackgroundAnomalous phantom visual perceptions coupled to an aversion and discomfort to some visual patterns (especially grating in mid-range spatial frequency) have been associated with the hyperresponsiveness in migraine patients. Previous literature has found fluctuations of alpha oscillation (8-14 Hz) over the visual cortex to be associated with the gating of the visual stream. In the current study, we examined whether alpha activity was differentially modulated in migraineurs in anticipation of an upcoming stimulus as well as post-stimulus periods.MethodsWe used EEG to examine the brain activity in a group of 28 migraineurs (17 with aura /11 without) and 29 non-migraineurs and compared their alpha power in the pre/post-stimulus period relative to the onset of stripped gratings.ResultsOverall, we found that migraineurs had significantly less alpha power prior to the onset of the stimulus relative to controls. Moreover, migraineurs had significantly greater post-stimulus alpha suppression (i.e event-related desynchronization) induced by the grating in 3 cycles per degree at the 2nd half of the experiment.ConclusionsThese findings, taken together, provide strong support for the presence of the hyperresponsiveness of the visual cortex of migraine sufferers. We speculate that it could be the consequence of impaired perceptual learning driven by the dysfunction of GABAergic inhibitory mechanism.

Error-related Alpha Suppression: Scalp Topography and (Lack of) Modulation by Modafinil.

Errors in performance trigger cognitive and neural changes that are implemented to adaptively adjust to fluctuating demands. Error-related alpha suppression (ERAS)-which refers to decreased power in the alpha frequency band after an incorrect response-is thought to reflect cognitive arousal after errors. Much of this work has been correlational, however, and there are no direct investigations into its pharmacological sensitivity. In Study 1 (n = 61), we evaluated the presence and scalp distribution of ERAS in a novel flanker task specifically developed for cross-species assessments. Using this same task in Study 2 (n = 26), which had a placebo-controlled within-subject design, we evaluated the sensitivity of ERAS to placebo (0 mg), low (100 mg), and high (200 mg) doses of modafinil, a wakefulness promoting agent. Consistent with previous work, ERAS was maximal at parieto-occipital recording sites in both studies. In Study 2, modafinil did not have strong effects on ERAS (a significant Accuracy × Dose interaction emerged, but drug-placebo differences did not reach statistical significance after correction for multiple comparisons and was absent after controlling for accuracy rate). ERAS was correlated with accuracy rates in both studies. Thus, modafinil did not impact ERAS as hypothesized, and findings indicate ERAS may reflect an orienting response to infrequent events.

Neural correlates of familiar and unfamiliar action in infancy

Behavioral evidence shows that experience with an action shapes action perception. Neural mirroring has been suggested as a mechanism underlying this behavioral phenomenon. Suppression of electroencephalogram (EEG) power in the mu frequency band, an index of motor activation, typically reflects neural mirroring. However, contradictory findings exist regarding the association between mu suppression and motor familiarity in infant EEG studies. In this study, we investigated the neural underpinnings reflecting the role of familiarity in action perception. We measured neural processing of familiar (grasp) and novel (tool-use) actions in 9- and 12-month-old infants. Specifically, we measured infants’ distinct motor/visual activity and explored functional connectivity associated with these processes. Mu suppression was stronger for grasping than for tool use, whereas significant mu and occipital alpha (indexing visual activity) suppression were evident for both actions. Interestingly, selective motor–visual functional connectivity was found during observation of familiar action, a pattern not observed for novel action. Thus, the neural correlates of perception of familiar actions may be best understood in terms of a functional neural network rather than isolated regional activity. Our findings provide novel insights on analytic approaches for identifying motor-specific neural activity while also considering neural networks involved in observing motorically familiar versus unfamiliar actions.

Does predictive cueing of presentation time modulate alpha power and facilitate visual working memory performance in younger and older adults?

Selective attention and working memory (WM) are vulnerable to age-related decline. Older adults perform worse on, and are less able to modulate alpha power (8–12 Hz) than younger adults in tasks involving cues about ‘where’ or ‘when’ a memory set will appear. However, no study has investigated whether alpha power is modulated by cues predicting the presentation time of a memory set. Here, we recorded electroencephalography while 24 younger (18–33 years) and 23 older (60–77 years) adults completed a modified delay match-to-sample task where participants were cued to the duration of a memory set (0.1 s or 0.5 s). We found: (1) predictive cues increased WM storage; (2) no differences in preparatory alpha power between predictive and neutral cue types, but preparatory alpha suppression was weaker in older adults; (3) retention period oscillatory power differed between presentation times, but these differences were no longer present when comparing trial types from the onset of the memory set; and (4) oscillatory power in the preparatory and retention periods were unrelated to performance. Our results suggest that preparatory alpha power is not modulated by predictive cues towards presentation time, however, reductions in alpha/beta power during visual WM retention may be linked to encoding, rather than retention.

Dynamics of alpha suppression and enhancement may be related to resource competition in cross-modal cortical regions

In the face of multiple sensory streams, there may be competition for processing resources in multimodal cortical areas devoted to establishing representations. In such cases, alpha oscillations may serve to maintain the relevant representations and protect them from interference, whereas theta band activity may facilitate their updating when needed. It can be hypothesized that these oscillations would differ in response to an auditory stimulus when the eyes are open or closed, as intermodal resource competition may be more prominent in the former than in the latter case. Across two studies we investigated the role of alpha and theta power in multimodal competition using an auditory task with the eyes open and closed, respectively enabling and disabling visual processing in parallel with the incoming auditory stream. In a passive listening task (Study 1a), we found alpha suppression following a pip tone with both eyes open and closed, but subsequent alpha enhancement only with closed eyes. We replicated this eyes-closed alpha enhancement in an independent sample (Study 1b). In an active auditory oddball task (Study 2), we again observed the eyes open/eyes closed alpha pattern found in Study 1 and also demonstrated that the more attentionally demanding oddball trials elicited the largest oscillatory effects. Theta power did not interact with eye status in either study. We propose a hypothesis to account for the findings in which alpha may be endemic to multimodal cortical areas in addition to visual ones.

Frontal Alpha Asymmetry in Children with Trauma Exposure.

The purpose of the current study was to investigate differences in frontal alpha asymmetry (FAA) between children (5-17 years) with or without histories of trauma exposure. EEG data were obtained from 165 children who participated in the Healthy Brain Network Initiative during rest with eyes open and closed. FAA during resting-state electroencephalography was significantly more negative in the trauma-exposed group, suggesting greater left lateralized FAA and avoidance-oriented motivation. Moreover, alpha suppression (difference in alpha amplitude between eyes open and eyes closed conditions) was marginally greater in the trauma-exposed group. The results suggest that early exposure to trauma may be associated with trait-level avoidance of environmental stimuli, which ultimately may be predictive of psychopathology, including posttraumatic stress disorder (PTSD). Study findings thus provide preliminary evidence of brain-based mechanisms that may confer risk for PTSD in the wake of early trauma exposure.

Maternal diabetes-mediated RORA suppression in mice contributes to autism-like offspring through inhibition of aromatase

Retinoic acid-related orphan receptor alpha (RORA) suppression is associated with autism spectrum disorder (ASD) development, although the mechanism remains unclear. In this study, we aim to investigate the potential effect and mechanisms of RORA suppression on autism-like behavior (ALB) through maternal diabetes-mediated mouse model. Our in vitro study in human neural progenitor cells shows that transient hyperglycemia induces persistent RORA suppression through oxidative stress-mediated epigenetic modifications and subsequent dissociation of octamer-binding transcription factor 3/4 from the RORA promoter, subsequently suppressing the expression of aromatase and superoxide dismutase 2. The in vivo mouse study shows that prenatal RORA deficiency in neuron-specific RORA null mice mimics maternal diabetes-mediated ALB; postnatal RORA expression in the amygdala ameliorates, while postnatal RORA knockdown mimics, maternal diabetes-mediated ALB in offspring. In addition, RORA mRNA levels in peripheral blood mononuclear cells decrease to 34.2% in ASD patients (n = 121) compared to the typically developing group (n = 118), and the related Receiver Operating Characteristic curve shows good sensitivity and specificity with a calculated 84.1% of Area Under the Curve for ASD diagnosis. We conclude that maternal diabetes contributes to ALB in offspring through suppression of RORA and aromatase, RORA expression in PBMC could be a potential marker for ASD screening.

Respiration aligns perception with neural excitability.

Recent studies from the field of interoception have highlighted the link between bodily and neural rhythms during action, perception, and cognition. The mechanisms underlying functional body-brain coupling, however, are poorly understood, as are the ways in which they modulate behavior. We acquired respiration and human magnetoencephalography data from a near-threshold spatial detection task to investigate the trivariate relationship between respiration, neural excitability, and performance. Respiration was found to significantly modulate perceptual sensitivity as well as posterior alpha power (8-13 Hz), a well-established proxy of cortical excitability. In turn, alpha suppression prior to detected versus undetected targets underscored the behavioral benefits of heightened excitability. Notably, respiration-locked excitability changes were maximized at a respiration phase lag of around -30° and thus temporally preceded performance changes. In line with interoceptive inference accounts, these results suggest that respiration actively aligns sampling of sensory information with transient cycles of heightened excitability to facilitate performance.

Alpha suppression indexes a spotlight of visual-spatial attention that can shine on both perceptual and memory representations.

Although researchers have been recording the human electroencephalogram (EEG) for almost a century, we still do not completely understand what cognitive processes are measured by the activity of different frequency bands. The 8- to 12-Hz activity in the alpha band has long been a focus of this research, but our understanding of its links to cognitive mechanisms has been rapidly evolving recently. Here, we review and discuss the existing evidence for two competing perspectives about alpha activity. One view proposes that the suppression of alpha-band power following the onset of a stimulus array measures attentional selection. The competing view is that this same activity measures the buffering of the task-relevant representations in working memory. We conclude that alpha-band activity following the presentation of stimuli appears to be due to the operation of an attentional selection mechanism, with characteristics that mirror the classic views of attention as selecting both perceptual inputs and representations already stored in memory.

Increased alpha suppression with age during involuntary memory retrieval.

Recent work suggests that while voluntary episodic memory declines with age, involuntary episodic memory, which comes to mind spontaneously without intention, remains relatively intact. However, the neurophysiology underlying these differences has yet to be established. The current study used electroencephalography (EEG) to investigate voluntary and involuntary retrieval in older and younger adults. Participants first encoded sounds, half of which were paired with pictures, the other half unpaired. EEG was then recorded as they listened to the sounds, with participants in the involuntary group performing a sound localization cover task, and those in the voluntary group additionally attempting to recall the associated pictures. Participants later reported which sounds brought the paired picture to mind during the localization task. Reaction times on the localization task were slower for voluntary than involuntary retrieval and for paired than unpaired sounds, possibly reflecting increased attentional demands of voluntary retrieval and interference from reactivation of the associated pictures respectively. For the EEG analyses, young adults showed greater alpha event-related desynchronization (ERD) during voluntary than involuntary retrieval at frontal and occipital sites, while older adults showed pronounced alpha ERD regardless of intention. Additionally, older adults showed greater ERD for paired than unpaired sounds at occipital sites, likely reflecting visual reactivation of the associated pictures. Young adults did not show this alpha ERD memory effect. Taken together, these data suggest that involuntary memory is largely preserved with age, but this may be due to older adults' greater recruitment of top-down control even when demand for such control is limited.