Psychophysiology

Successful behavioral adaptation requires an ongoing assessment of rewarding outcomes based on ones current state. A frontocentral ERP associated with reward feedback, the reward positivity (RewP), has been linked to reflect information about reward value and motivational states. It is, however, unclear if changes in the RewP are influenced by changes in reward value as a function of motivational state. To examine this, hungry participants (n=31) completed two rounds of a modified Doors Task incorporating Pavlovian conditioning during EEG recordings and obtained feedback associated with sweet and savory food reinforcers equally matched in pleasantness and desirability. Participants underwent reinforcer devaluation, a paradigm designed to isolate inference-based behavior based on decreasing reward value, in between rounds by eating one of the foods to satiety. Prior to devaluation, participants were hungry and rated both food reinforcers equally pleasant. After devaluation, participants were sated and rated the devalued food, but not the non-devalued food, significantly less pleasant, suggesting a sensory-specific change in reward value. Logistic regression of win-stay/lose-switch behavior during the Doors Task shows participants made sensory-specific adjustments in food preferences during post-devaluation. Non-parametric permutation tests based on the tmax statistic performed revealed no significant differences in RewP amplitudes, suggesting the RewP is insensitive to reinforcer devaluation. This could not be explained by differences in perceived pleasantness or desirability. These findings suggest that affective and motivational factors such as tracking inferences based on decreases in reward value did not modulate the RewP.

Motivated by the Neurovisceral Integration Model (NVI) of cardiac vagal control, we investigated the relationship between relative left frontal activity (rLFA) and vagally mediated heart rate variability or respiratory sinus arrhythmia (RSA) in 287 participants, half of whom had a history of depression. We hypothesized that there would be a within-person association of rLFA and RSA such that when RSA is lower rLFA would also be lower (Hypothesis I). Moreover, it was hypothesized that this within-subject association would be moderated by a history of depression (Hypothesis II). Metrics of rLFA and RSA were derived from concurrent electroencephalogram and electrocardiogram recordings. The logarithmic difference in EEG alpha power between the homologous right and left electrodes (Ln (Right/Left)) in the frontal region was used to index rLFA. A Hilbert transform was applied to the mean-centered and bandpass-filtered (0.12-.40 Hz) inter-beat interval (IBI) time series to get a fine-grained measure (in the time domain) of RSA. A linear mixed ANOVA model with rLFA as the dependent variable and RSA as the main fixed effect found that participants had less rLFA during epochs when they had lower RSA, which was consistent with the prediction from Hypothesis I. Contrary to the prediction from Hypothesis II, the within-person association of RSA and rLFA was not moderated by a history of depression. However, the association between RSA and rLFA varied across the four pairs of frontal electrodes that we examined. Thus, more research is needed to determine the spatial extent of this association, e.g., examining the relationship between source-localized rLFA and RSA.

To effectively function in an ever-changing environment, the brain is proposed to make predictions about upcoming information. However, the association between prediction and memory formation and the role of between-subject neural variability in this relationship is unclear. To shed light on the relationship between prediction and memory, the present study reanalysed data from Jano, Chatburn, and colleagues (2024). In the original experiment, participants were exposed to naturalistic images in predictable and unpredictable four-item sequences, after which their memory was tested using an old/new paradigm. In the present analysis (N = 46), N400 amplitude and oscillatory power during learning was measured to gauge processes related to prediction error and memory encoding, respectively. This activity was compared with subsequent memory outcomes and individual alpha frequency (IAF) calculated at rest. Linear mixed-effects regressions revealed an alpha power subsequent memory effect that was not related to the amplitude of the N400, suggesting that memory encoding may occur independently of the level of prediction error. Notably, IAF influenced the relationship between theta power, N400 amplitude and subsequent memory, implying that the electrophysiological conditions for successful memory formation differ between individuals. Consequently, the degree to which prediction errors (presumably gauged via the N400) drive memory encoding could depend on inter-individual variability in intrinsic neural activity. These findings emphasise the flexible nature of memory, whilst having potential implications for prediction error-driven accounts of learning.

Numerous studies have demonstrated rapid (< 100 ms) visuo-cortical differentiation of threat-associated faces. This may be due to low-spatial frequency (LSF) visual information originating from magnocellular pathways. Yet it remains unclear whether potentially magnocellular fear signals extend beyond evolutionarily prepared emotional faces and whether they are subject to short-term neuroplasticity. If so, spatial frequency characteristics should modulate processing of faces with newly acquired threat-relevance. Furthermore, it is unknown whether sub-bands of the visual spectrum are associated with autonomic arousal. Using a differential fear-conditioning paradigm, this study tested whether early visual attentional capture, indicated by the P1 event-related potential component, prioritizes LSF information of threat-associated faces with neutral expressions. Additionally, it was tested whether such effects would be paralleled by threat differentiation in the skin conductance response (SCR). For contingency aware participants, stimulus ratings confirmed successful fear conditioning and participants showed a selective left-hemispheric enhancement of the P1 in response to LSF threat-faces. By contrast, CS differentiation in the SCR was not modulated by spatial frequencies but by stimulus duration, with longer CS presentations resulting in larger SCR to threat compared to neutral faces. For contingency unaware participants, trial-by-trial amplitudes of P1 and SCR were positively correlated. Data support the notion that magnocellular-cortical pathways adapt quickly to novel threat-associations and facilitate rapid threat retrieval even for perceptually neutral faces. However, at least in the short term, these signals do not necessarily associate with anticipatory arousal in SCR. Impact statementOur electroencephalography (EEG) study provides evidence for distinct contributions of subcortical signals during early visual perception of fear-conditioned faces (P1 event-related potential) but not autonomic arousal (skin conductance response). Instead, skin conductance responses reflected conscious anticipatory arousal irrespective of the visual pathway. Together, these results reveal parallel but dissociable mechanisms of fear perception that are differentially sensitive to visual properties of threat-associated faces.

A proactive mechanism has been postulated to promote successful inhibition (Kenemans, 2015). Specifically, this mechanism is thought to operate before any action demanding or countermanding event has occurred. In the current study, we investigated whether EEG theta power could reflect this mechanism, in a sample of healthy individuals performing a stop-signal paradigm. By comparing frontal theta power preceding failed versus successful stop trials, we tested whether frontal theta is predictive of inhibition success. We hypothesized that proactive cognitive control manifests in frontal theta power preceding a countermanding go-stop event. Our results demonstrate that frontal theta is indeed higher preceding successful as compared to preceding failed stopping events. We also show that frontal theta power preceding stopping events is associated with Stop-Signal Reaction Times (SSRT), with a higher theta being indicative of shorter SSRTs. This association was not present for go-RT. This study may be the first to reveal a relationship between lower frontal theta power and subsequent stopping failure, suggesting thetas role in proactive response inhibition.

We report the first use of ERP measures to identify text engagement differences when reading digitally or in print. Depth of semantic encoding is key for reading comprehension, and we predicted that deeper reading of expository texts would facilitate stronger associations with subsequently-presented related words, resulting in enhanced N400 responses to unrelated probe words and a graded attenuation of the N400 to related and moderately related words. In contrast, shallow reading would produce weaker associations between probe words and text passages, resulting in enhanced N400 responses to both moderately related and unrelated words, and an attenuated response to related words. Behavioral research has shown deeper semantic encoding of text from paper than from a screen. Hence, we predicted that the N400 would index deeper reading of text passages that were presented in print, and shallower reading of texts presented digitally. Middle-school students (n = 59) read passages in digital and print formats and high-density EEG was recorded while participants completed single-word semantic judgment tasks after each passage. Following digital text reading, the N400 response pattern anticipated for shallow reading was observed. Following print reading, the N400 response pattern expected for deeper reading was observed for related and unrelated words, although mean amplitude differences between related and moderately related probe words did not reach significance. These findings provide evidence of differences in brain responses to texts presented in print and digital media, including deeper semantic encoding for print than digital texts.

Recent studies have highlighted the essential role of interoception in healthy emotional processing and the pathology of major depressive disorder. However, it is unclear how individual differences in healthy people with high depression risk (HDR; i.e., individual differences in depression risk) are related to the neurophysiological underpinnings of interoception and emotional reactions under different degrees of certainty. We examined whether an individuals depression risk mediates the relationships of heartbeat-evoked potentials (HEPs), a neurophysiological marker of cardiac interoception, with heart rate (HR) and heart rate variability (HRV), indices for cardiac reactivity. In a concurrent electroencephalogram-electrocardiogram (EEG-ECG) experiment, 26 healthy participants completed an emotion-evoking picture-evaluation task. Each trial began with a differential auditory cue that was associated with the certainty of subsequently seeing a pleasant or unpleasant picture. The results showed the following: after participants saw a cue of uncertainty, HDR was associated with 1) reduced neural activity when anticipating upcoming pictures and 2) enhanced physiological reactions to unexpected, unpleasant pictures. These results suggest that weaker predictions and augmented prediction errors for negative emotional stimuli reflect depression risk. Moreover, depression risk significantly mediated the relationships between HEPs and HR and between HEPs and HRV for unexpected, unpleasant stimuli. This study provides evidence that interoception and autonomic cardiac regulation may be altered by depression risk. This highlights the insights provided by specific indices of brain-heart interactions, such as HEPs, into the underlying activity of the autonomic nervous system and unique interoceptive disturbances associated with depression risk.

Behavioral evidence shows that anxious individuals tend to be distracted by irrelevant stimulation not only for threat-related stimuli but also for non-emotional neutral stimuli. These findings suggest that anxious individuals may have a general impairment of attentional control, especially inhibition function. However, the neural mechanism underlying the anxiety-related impairment in attentional control is unclear. Here, in a visual search task with geometric stimuli, we examined attentional processing of the non-emotional neutral distractor on participants with different levels of anxiety, using the event-related-potential (ERP) indices of attentional selection (N2 posterior contralateral [N2pc]) and top-down inhibition (distractor positivity [Pd]). We found that distractor-evoked Pd amplitudes were negatively correlated with trait-anxiety scores, i.e., the higher the level of anxiety, the worse the ability of attentional inhibition. In contrast, the amplitudes of distractor-evoked N2pc did not vary with anxiety levels, suggesting that trait-anxiety level does not affect stimulus-driven attentional capture. We also observed attentional processing of target stimuli and found that the peak latency of target-evoked N2pc was delayed as anxiety levels rise, suggesting that anxiety impairs the efficiency of top-down attentional selection of the target. The present study provides direct neurophysiological evidence for general anxiety-related impairment of attentional control.

The error-related negativity (ERN) is a neural correlate of error monitoring often used to investigate individual differences in developmental, mental health, and adaptive contexts. However, limited experimental control over errors presents several confounds to its measurement. An experimentally controlled disturbance to standing balance evokes the balance N1, which we previously suggested may share underlying mechanisms with the ERN based on a number of shared features and factors. We now measure whether the balance N1 and ERN are correlated across individuals within two small groups (N=21 young adults and N=20 older adults). ERNs were measured in arrow flanker tasks using hand and foot response modalities (ERN-hand and ERN-foot). The balance N1 was evoked by sudden slip-like movements of the floor while standing. The ERNs and the balance N1 showed good and excellent internal consistency, respectively, and were correlated in amplitude in both groups. One principal component strongly loaded on all three evoked potentials, suggesting that the majority of individual differences are shared across the three ERPs. However, there remains a significant component of variance shared between the ERN-hand and ERN-foot beyond what they share with the balance N1. It is unclear whether this component of variance is specific to the arrow flanker task, or something fundamentally related to error processing that is not evoked by a sudden balance disturbance. If the balance N1 were to reflect error processing mechanisms indexed by the ERN, balance paradigms offer several advantages in terms of experimental control over errors.

The arousal-biased competition theory posits that inducing arousal increases attentional priority of salient stimuli while reducing priority of non-pertinent stimuli. However, unlike in young adults, older adults rarely exhibit shifts in priority under increased arousal, and prior studies have proposed different neural mechanisms to explain how arousal differentially modulates selective attention in older adults. Therefore, we investigated how the threat of unpredictable shock differentially modulates attentional control mechanisms in young and older adults by observing eye movements. Participants completed two oculomotor search tasks in which the salient distractor was typically captured by attention (singleton search) or proactively suppressed (feature search). We found that arousal did not modulate attentional priority for any stimulus among older adults nor affect the speed of attention processing in either age group. Furthermore, we observed that arousal modulated pupil sizes and found a correlation between evoked pupil responses and oculomotor function. Our findings suggest age differences in how the locus coeruleus-noradrenaline system interacts with neural networks of attention and oculomotor function. Highlights- Increased arousal modulates attention saliency priority but not processing speed - Older adults do not exhibit shifts in stimulus priority under elevated arousal - Proactive suppression of salient stimuli persists even during increased arousal - Threat of unpredictable shock increases pupil sizes and decreases evoked responses - Eye movements may be able to help assess locus coeruleus function

Mindfulness meditation involves training attention, commonly towards the current sensory experience, with an attitude of non-judgemental awareness. Theoretical perspectives suggest meditation alters the brains predictive processing mechanisms, increasing the synaptic gain and precision with which sensory information is processed, and reducing the generation or elaboration of higher-order beliefs. Recent research suggests that forwards and backwards travelling cortical alpha waves provide an indication of these predictive processing functions. Here, we used electroencephalography (EEG) to test whether the strength of forwards and backwards travelling cortical alpha waves differed between experienced meditators and a matched sample of non-meditators, both during an eyes-closed resting state (N = 97) and during a visual cognitive (Go/No-go) task (N = 126). Our results showed that meditators produced stronger forwards travelling cortical alpha waves compared to non-meditators, both while resting with their eyes closed and during task performance. Meditators also exhibited weaker backwards travelling cortical alpha wave strength while resting with their eyes closed. These results may be indicative of a neural mechanism underpinning enhanced attention associated with meditation practice, as well as a potential neural marker of the reductions in resting mind-wandering that are suggested to be associated with meditation practice. The results also support models of brain function that suggest attention modification can be achieved by mental training aimed at increased processing of sensory information, which might be indexed by greater strength of forwards travelling cortical alpha waves.

Age-related changes to the power and frequency of the brains oscillatory activity have been reported by an extensive literature. In contrast, the influence of advancing age on the shape of oscillation waveforms, a characteristic with increasingly recognised physiological and functional relevance, has not been previously investigated. To address this, we examined the shape of alpha and beta band oscillations from electroencephalography (EEG) data recorded during performance of simple and go/no-go reaction time tasks in 33 young (23.3 {+/-} 2.9 years, 27 females) and 27 older (60.0 {+/-} 5.2 years, 23 females) adults. The shape of individual cycles was characterised using instantaneous frequency, and then decomposed into waveform motifs using principal component analysis. This analysis identified four principal components (one from the alpha band, 3 from the beta band) that were uniquely influenced by the different motor tasks and/or age. These each described different dimensions of shape and tended to be modulated during the reaction phase of each task. However, the way in which each facet of shape varied during the task was unrelated to motor performance, indexed via reaction time, in either group or band. Our results suggest that although oscillation shape is task-dependent, the nature of this effect is altered by advancing age. While these outcomes demonstrate the utility of this approach for understanding the neurophysiological effects of ageing, future work that more clearly links these outcomes with function will be critical.

Affective experiences are inevitably accompanied by physiological changes, however it is still a matter of intense debate whether events evoking similar affective experiences produce comparable physiological responses (fingerprint hypothesis) or variation is the norm within individuals (population hypothesis). We reanalyzed data from two independent samples (N = 491; N = 64), using representational similarity analysis (RSA) to examine the trial-by-trial similarity patterns of subjective experience of valence and arousal and affect-related physiological measures (skin conductance [SCR] and startle blink responses). Across different affect-inducing (passive picture viewing, passive sound listening, imagery tasks) tasks and samples (Ns=491, 64), we observed strong-to-decisive evidence for a correspondence between SCR and startle responses and models of arousal and valence that assume variation, especially between trials generally evoking higher responses. Our results show that similar affective experiences are rather reflected by distinct physiological responses and emphasize the importance of considering intraindividual variability in future studies to better understand how physiological changes contribute to conscious affective experiences in humans.

Aversive conditioning prompts reliable changes in the power of EEG alpha-band oscillations and pupil dilation. Both variables have been used to test hypotheses on the acquisition, generalization, and extinction of conditioned threat. Existing studies have largely relied on trial averages and group-level analyses. Thus, the variability of these physiological markers to aversive learning at the subject level is currently unknown. Comparisons of group-level analyses in prior studies suggest that pupil dilation and EEG-alpha activity capture complementary information. However, to date, no study has directly compared these two markers in terms of their effect sizes at the level of individual participants. The present study employed Bayesian multilevel modeling to quantify the variability of conditioning effect estimates for alpha-band power and pupillometry. Estimates were examined at the group level and at the participant level, across two conditioning paradigms, involving visual and auditory cues. Although the two metrics shared similar effect sizes at the group level, participant-level variability in these effect sizes was substantially higher for pupil-dilation compared to alpha-power, and this finding was replicated across both paradigms. These findings have important implications for clinical and inter-individual difference research which requires both the quantification of effects at the participant-level as well as meaningful variability between-participants that can be linked to relevant differences such as anxiety.

Previous research has demonstrated that negative emotional faces dilate time perception, however, the mechanisms underlying this phenomenon are not fully understood. Previous attempts focus on the pacemaker-accumulator model of time perception, which includes a clock, memory, and decision-making stage, wherein emotion affects one of these stages; possibly by increasing pacemaker rate via arousal, increasing accumulation rate via attention, or by biasing decision-making. To further investigate the stage(s) that emotion is affecting time perception we conducted a visual temporal bisection task with sub-second intervals while recording 64-channel electroencephalogram (EEG). To separate the influence of face and timing responses the temporal stimulus was preceded and followed by a face stimulus displaying a neutral or negative expression creating three trial-types: Neg[-&gt;]Neut, Neut[-&gt;]Neg, or Neut[-&gt;]Neut. The data revealed a leftward shift in bisection point (BP) in Neg[-&gt;]Neut and Neut[-&gt;]Neg suggesting an overestimation of time. Neurally, we found the face-responsive N170 component was larger for negative faces and the N1 and contingent negative variation (CNV) were larger when preceded by a negative face. We also found an interaction effect between condition and response for the late positive component of timing (LPCt) and a significant difference between response (short/long) in the neutral condition. We conclude that a preceding negative face affects the clock stage leading to more pulses being accumulated, either through attention or arousal, as indexed by a larger N1, CNV, and N170; whereas viewing the negative face second biased decision-making leading to "short" responses being less likely, as evidenced by the LPCt.

The choice of sampling rate is a critical preprocessing step in event-related potential (ERP) research, yet its impact on different analytic approaches remains underexplored. In this study, we systematically evaluated how downsampling affects data quality measured via Standardized Measurement Error (SME), conventional univariate ERP metrics (mean amplitude, peak amplitude, peak latency, and 50% area latency), and multivariate pattern analysis (MVPA; decoding). We analyzed seven commonly studied ERP components: P3, N400, N170, N2pc, mismatch negativity, error-related negativity, and lateralized readiness potential collected from neurotypical young adults. Results showed that both amplitude- and latency-based scores were significantly affected by sampling rate changes, particularly for 50% area latency that presented increased SME (reduced data quality) at lower sampling rates. Amplitude-based measures and their corresponding effect sizes were more influenced by sampling rate variations, whereas latency-based measures were comparatively stable across different temporal resolutions. In contrast, multivariate decoding performance remained highly robust, with decoding accuracy and effect sizes showing minimal variation even at the lowest sampling rate (e.g., 64 Hz). Overall, these findings suggest that caution should be exercised when employing lower sampling rates for data quality and conventional univariate ERP analyses. Nevertheless, lower sampling rates (e.g., 64 Hz) may still be appropriate for decoding analyses, particularly in studies where precise temporal resolution is not critical. For researchers analyzing ERP data with similar components, noise levels, and participant populations as in this study, following these recommendations should yield robust statistical power.

The error-related negativity (ERN) and correct-related negativity (CRN) are event-related potentials (ERPs) that reflect performance monitoring following error and correct responses, respectively. Prior work demonstrates the ERN is sensitive to the motivational significance of errors, which increases under social observation. However, most studies testing how social observation impacts performance monitoring rely on trial-averaged ERPs, potentially obscuring meaningful fluctuations in ERN/CRN over time. Here, we had participants complete a Flanker task twice (social observation vs. alone) and employed mixed-effects modeling of single-trial ERPs to test if social observation impacts ERN/CRN trajectories over short (within blocks) or long (between blocks) timescales. We found that social observation selectively influenced ERN/CRN trajectories over short timescales: for blocks performed under social observation (but not alone), ERN magnitudes increased across trials and CRN magnitudes decreased. At longer timescales, ERN/CRN significantly decreased across all blocks, regardless of social observation and consistent with a vigilance decrement. To our knowledge, this is the first demonstration that social observation influences performance monitoring trajectories over short timescales. Results highlight the importance of analyzing ERN/CRN trajectories over relatively short timescales to fully characterize the impact of social observation on performance monitoring dynamics. These findings lay the groundwork for future investigation into whether social observation interacts with individual differences in motivation/affect to differentially impact performance monitoring dynamics.

Motion-Position Illusions (MPIs) involve the position of an object being misperceived in the context of motion (i.e. when the object contains motion, is surrounded by motion, or is moving). A popular MPI is the flash-lag effect, where a static object briefly presented in spatiotemporal alignment with a moving object, is perceived in a position behind the moving object. Recently, Cottier et al. (2023) observed that there are stable individual differences in the magnitude of these illusions, and possibly even their direction. To investigate the possible neural correlates of these individual differences, the present study explored whether a trait-like component of brain activity, individual alpha frequency (IAF), could predict individual illusion magnitude. Previous reports have found some correlations between IAF and perceptual tasks. Participants (N=61) viewed the flash-lag effect (motion and luminance), Frohlich effect, flash-drag effect, flash-grab effect, motion-induced position shift, twinkle-goes effect, and the flash-jump effect. In a separate session, five minutes of eyes-open and eyes-closed resting state EEG data was recorded. Correlation analyses revealed no evidence for a correlation between IAF and the magnitude of any MPIs. Overall, these results suggest that IAF does not represent a mechanism underlying MPIs, and that no single shared mechanism underlies these effects. This suggests that discrete sampling at alpha frequency is unlikely to be responsible for any of these illusions.

Research using electroencephalography (EEG) has shown that individual differences in visual working memory capacity are related to slow-wave event-related potentials (ERPs) and suppression of alpha-band oscillatory power during the delay period of memory tasks. However, recent evidence suggests that changes in non-oscillatory (aperiodic) features of the EEG signal are related to working memory performance. We assessed several features of task-related changes in aperiodic activity including its spatial distribution, the effect of memory load, and the relationships between aperiodic activity, memory capacity, slow-wave ERPs, and alpha suppression. Eighty-four healthy individuals performed a continuous recall working memory (WM) task consisting of 2, 4 or 6 coloured squares while EEG was recorded. Aperiodic activity during a baseline and WM delay period was quantified by fitting a model to the background of the EEG power spectra, which returned parameters describing the slope (exponent) and broadband offset of the spectra. The aperiodic exponent decreased (i.e., slope flattened) in lateral parieto-occipital electrodes but increased (i.e., slope steepened) in fronto-central electrodes during the WM delay period, whereas the offset decreased over parieto-occipital electrodes. These task-related changes in aperiodic activity did not differ between memory loads. Larger increases in the aperiodic exponent were associated with higher working memory capacity measured from both the WM task and a separate battery of complex span tasks, a relationship that was independent of slow-wave ERPs and alpha suppression. Our findings suggest that WM task-related changes in aperiodic activity are region specific and reflect an independent neural mechanism that is important for general working memory ability.

Adolescence is a stage of development characterized by neurodevelopmental specialization of cognitive processes. In particular, working memory continues to improve through adolescence, with increases in response accuracy and decreases in response latency continuing well into the twenties. Human electroencephalogram (EEG) studies indicate that gamma oscillations (35-65 Hz) during the working memory delay period support the maintenance of mnemonic information guiding subsequent goal-driven behavior, which decrease in power with development. Importantly, recent electrophysiological studies have shown that gamma events, more so than sustained activity, may underlie working memory maintenance during the delay period. However, developmental differences in gamma events during working memory have not been studied. Here, we used EEG in conjunction with a novel spectral event processing approach to investigate age-related differences in transient gamma band activity during a memory guided saccade (MGS) task in 164 10- to 30-year-olds. Total gamma power was found to significantly decrease through adolescence, replicating prior findings. Results from the spectral event pipeline showed age-related decreases in the mean power of gamma events and trial-by-trial power variability across both the delay period and fixation epochs of the MGS task. In addition, we found that while event number decreased with age during the fixation period, it did not appear to change during the delay period resulting in an increasing difference between the number of events during fixation and delay period with development, suggesting that as working memory develops there is greater specificity for gamma events supporting working memory. While average power of the transient gamma events was found to mediate age-related changes in total gamma power, the number of gamma events was unrelated to total power, suggesting that the power of gamma events may underlie the sustained gamma activity seen in EEG literature while the number of events may directly support age-related improvements in working memory maintenance. Our findings provide compelling new evidence for mechanistic changes in neural processing characterized by refinements in neural function as behavior becomes optimized in adulthood.