Theta EEG dynamics of the error-related negativity

“These findings have led to the interpretation that the ERN is indicative of error monitoring (Falkenstein et al., 1991, 2000; Gehring et al., 1993) or response competition (Carter et al., 1998) processes.”

“Similarly, in the time-frequency domain, EEG studies showed Theta (4-7Hz) and Alpha (8-13Hz) synchronizations over fronto-central electrodes as a marker of the performance monitoring system activity (Luu et al., 2004; Trujillo and Allen, 2007; Cavanagh et al., 2009; Cohen, 2011).”

“The ERS is characterized by an increased oscillatory activity in the theta frequency band-pass region (4-7.5 Hz) occurring approximately in the same time window and spatial location as the Ne (Trujillo and Allen, 2007).”

“After, we applied another bandpass filter between 1 Hz and 10 Hz (Butterworth, fourth order, zero-phase) since numerous studies found the lower frequency bands to have an essential role in error processing [39,71,75].”

“The wavelet cycles varied from 3 to 8 in logarithmically spaced steps, to obtain an optimal trade-off between temporal and frequency resolutions (Trujillo and Allen, 2007).”

“…Goss, Coles, Meyer, & Donchin, 2018), peaks from 50 to 100 ms after an error, and mainly emerges from the anterior cingulate cortex (ACC; Dehaene, Posner, & Tucker, 1994; Debener et al., 2005; Luu, Tucker, & Makeig, 2004; Holroyd, Dien, & Coles, 1998; Miltner et al., 2003; Trujillo & Allen, 2007).”

“During various tasks, midfrontal theta power is increased around (cid:0) 100 to 500 ms relative to error commission (Kalfao ˘ glu et al., 2018; Novikov et al., 2015; Trujillo and Allen, 2007; Yordanova et al., 2004).”

“This 3-factor solution identified a clear MFC theta band factor maximal immediately following the response, consistent with prior work investigating post-error theta (Cavanagh et al., 2009), which has also been shown to contribute to the ERN (Luu et al., 2004; Trujillo and Allen, 2007).”

“We adjusted the number of wavelet cycles from 3 to 8 in logarithmically spaced steps to achieve an optimal balance between temporal and frequency precision (Trujillo and Allen 2007).”

“While an increase in Fm θ may occur following both correct and incorrect responses in highly demanding tasks, Fm θ increase is a far more robust phenomenon during performance errors (Luu and Tucker, 2001; Luu et al., 2004; Yordanova et al., 2004; Trujillo and Allen.”

“…et al., 2010; Yordanova et al., 2004), for theta 250 ms preresponse to 500 ms postresponse (Luu, Tucker, & Makeig, 2004; Mazaheri et al., 2009; Trujillo & Allen, 2007), and for low beta from 500 ms preresponse (Grent-'tJong, Oostenveld, Jensen, Medendorp, & Praamstra, 2013) to 850 ms…”

“EEG signatures of errors also differ from those of conflict in correct responses because of their additional recruitment of delta power (Yordanova et al., 2004; Cohen and van Gaal, 2014) and phase-locked component (Trujillo and Allen, 2007; Munneke et al., 2015).”

“…the other hand, theta-ERS has been associated with conflict-related processes such as conflict detection and post-error correction, reflecting reactive control (Cavanagh et al., 2009; Cavanagh and Frank, 2014; Cohen and Donner, 2013; Luu et al., 2004; Trujillo and Allen, 2007; Wei and Zhou, 2020).”

“These midfrontal theta oscillations are considered to reflect the computations needed for cognitive control, monitoring, error processing, and adjusting of temporally sequenced actions (Cavanagh and Frank 2014; Cohen 2014; Trujillo and Allen 2007; Valadez and Simons 2017; Womelsdorf et al. 2010).”

“…findings from the decision-making and cognitive control literature have revealed that both ERPs and theta power track errors and negative feedback signals prior to and following a response ( Cavanagh & Frank, 2014; Cavanagh et al., 2010; Luft, 2014; Luu et al., 2004; Trujillo & Allen, 2007).”

“measured at midline fronto-central electrodes in the range of its peak negativity between 50-100 milliseconds following error commission [11], [12].”

“Moreover, increased frontal theta band activity (TBA) is associated with better performance in conflict monitoring (Cavanagh et al., 2009; Eschmann et al., 2018; Trujillo and Allen, 2007) as well as response inhibition (Funderud et al., 2012; Kamarajan et al., 2004; Nigbur et al., 2011).”

“However, following errors, the increase in the non-phase-locked theta power was larger and longer-lasting than phase-locked theta power [23,66].”

“The typical electrophysiological pattern of feedback processing was observed on our probabilistic gambling task, consisting of a clear frontocentral FRN, greater parietal delta and frontal beta-gamma activities after gains, as compared to losses, and increased frontal theta activity after losses, as compared to after…”

“…2014; Wascher et al. 2014) but also anxious responses (e.g., Cavanagh and Shackman 2015; Osinsky et al. 2017), and feedback processing (e.g., Fu et al. 2023; Trujillo and Allen 2007; also see Rawls et al. 2020), which has often been reported to be distorted in MD patients (e.g., Keren et al. 2018).”

“…6.5 Hz, often observed at EEG electrodes FCz or occasionally Cz, with less occurrence in lateral or more anterior regions (Cohen & Cavanagh, 2011; Cohen & van Gaal, 2014; Hanslmayr et al., 2008; Luu et al., 2003, 2004; Past € otter et al., 2010; Swingler et al., 2017; Trujillo & Allen, 2007).”

“…and interareal processing mechanisms ( Fries, 2009, 2015; Wang, 2010), and modulations of oscillatory activity have been implicated in various cognitive functions like cognitive control, learning, memory, and action regulation ( Cavanagh & Frank, 2014; Herweg et al., 2020; Trujillo & Allen, 2007).”

“It is hypothesized that large-scale functional integration across different frequency bands supports flexible behavior adaption to improve the performance after an error and thus results in an increase of PAC following error response [22].”

“…& Yamamoto, 2010), as well as other control-related processes such as response error and feedback processing (Bernat, Nelson, Steele, Gehring, & Patrick, 2011; Cohen, Elger, & Ranganath, 2007; Gehring & Willoughby, 2004; Trujillo & Allen, 2007; Yordanova, Falkenstein, Hohnsbein, & Kolev, 2004).”

“A temporally similar negative potential, the error-related negativity (ERN), is observed in response-locked, event-related potentials (ERPs) in participants solving a choice reaction time task incorrectly (71 – 81).”

“The flicker frequency was set at 7 Hz to study the effects of using an SSVEP frequency in the theta band that overlaps with the ErrP spectral response 95–97 and the action time was kept the same as D3 (3 seconds).”

“Theta band oscillations are known to reflect error detection and correction process (Trujillo & Allen, 2007).”

“…(4–7 Hz) may be one mechanism that underlies functional communication between networks involving medial prefrontal cortex (mPFC) and lateral prefrontal cortex (lPFC) regions during the ERN (25–75 ms time window) [69], [71], [72], [73], all analysis was performed for this time and frequency range.”

“Indeed, prior data suggest that ERN is dominated by phase-locking of intermittent theta-band (Trujillo & Allen, 2007), but that the observation of an error also elicits non-phase locked activity; thus, not all mid-frontal activity is associated to oERN generation (Moreau et al., 2020; Pezzetta et…”

“…error-related response (Keil, Weisz, Paul-Jordanov, & Wienbruch, 2010; Luu, Tucker, & Makeig, 2004; Roger, Bénar, Vidal, Hasbroucq, & Burle, 2010; Trujillo & Allen, 2007; Yeung, Bogacz, Holroyd, Nieuwenhuis, & Cohen, 2007) and cognitive load and control (Cavanagh, Zambrano-Vazquez, & Allen,…”

“Theta has also been demonstrated to be enhanced during losses in gambling tasks (Leicht et al., 2013; Marco-Pallares et al., 2008) and flanker tasks designed to elicit errors (Trujillo & Allen, 2007).”

“The increased ERN negativity and elevated delta–theta power following false alarms replicate prior evidence that these neural markers are sensitive to error commission, particularly under conditions of inhibitory failure 33,36,46 .”

“Previous literature has linked changes in midfrontal theta to cognitive control 8,9 during conflict and error monitoring, 12,66 attentional effort, 34,67 expectancy violation, 7 and value-based decision-making.”

“Both FRN and the related theta responses are associated with sources in the anterior cingulate cortex (ACC) and the medial prefrontal cortex (Miltner et al. 1997; Holroyd et al. 1998; Dehaene et al. 2003; Trujillo and Allen 2007; Beldzik et al. 2022).”

“The ERN has been closely linked to frontal midline theta oscillations, which represent the engagement of cognitive control during error processing (Cavanagh et al. 2009; Keil et al. 2010; Luu et al. 2004; Trujillo and Allen 2007).”

“…with better performance on a variety of cognitive control tasks, including conflict monitoring (Cavanagh et al., 2009; Eschmann et al., 2018; Trujillo & Allen, 2007) and working memory updating (Itthipuripat et al., 2013; Jacobs et al., 2006; Jensen & Tesche, 2002; Klimesch et al., 2006).”

“We adjusted the number of wavelet cycles from 3 to 8 in logarithmically spaced steps to achieve an optimal balance between temporal and frequency precision (Trujillo and Allen, 2007).”

“Theta oscillations exhibit maximal increases in power (magnitude) and synchrony (consistent phase alignment) over MFC following error responses (Buz-zell et al., 2019; Cavanagh et al., 2009; Luu et al., 2004; Trujillo & Allen, 2007).”

“Given that previous studies have described an association between the FRN and medial-frontal theta oscillatory activity (4–8 Hz) (Cavanagh et al., 2010; Cohen et al., 2007; Marco-Pallarés et al., 2008; Padrão et al., 2013; Trujillo and Allen, 2007) we performed waveletbased time frequency analysis of the EEG data to s…”

“…(Abdul Rahman et al., 2017; Brooker et al., 2020; Cragg et al., 2009; Hoyniak, 2017; Johnstone et al., 2007; Jonkman, 2006; Kim et al., 2007; Lamm et al., 2006; Lo, 2018; Torpey et al., 2012; Trujillo & Allen, 2007; Wiersema et al., 2007), all variables were estimated on a frontal ROI (F3/F4).”

“Data preprocessing included temporal filtering be-tween 1 and 10 Hz, a frequency range of particular interest in error processing (e.g., Trujillo et al. [47]), which also attenuates excessive contamination from muscle artifacts.”

“Frontal-midline theta activity reflects dynamic coordination among these regions and provides a temporally sensitive index of prediction–outcome evaluation [ 17, 18 ].”

“Theta activity is also related to processing error/conflict monitoring (Trujillo and Allen 2007 ), self-regulation (Ertl et al. 2013 ; Knyazev 2007 ), stress, and anxiety (Sreekrishnan et al. 2014 ).”

“Induced (non-phase-locked) spectral power reflecting oscillatory activity without contribution of evoked response was obtained (Luu et al., 2004; Trujillo & Allen, 2007).”

“The negative correlation spanning from posterior to central regions may re fl ect engagement of medial frontal and central neural generators involved in error processing, which are well documented in the ERP literature (Trujillo and Allen, 2007; Wessel, 2012).”

“The error-related negativity has been reported as a phase resetting of Theta (Trujillo and Allen 2007).”

“Theta is observed during surprising events, such as negative feedback (Cavanagh, Frank, Klein, & Allen, 2010; Trujillo & Allen, 2007; van de Vijver et al., 2011; Andreou et al., 2017; Brincat & Miller, 2015; Loonis et al., 2017) and stimulus conflict (Muralidharan et al., 2023; Choi et al., 2024).”

“Moreover, FM-theta activity has been linked directly to ERN peak amplitude (Luu et al. 2004; Trujillo and Allen 2007).”

“They noted increased N2 amplitude, an event-related potential (ERP) component thought to be driven by phase-locked FMT (Cavanagh et al., 2009; Luu et al., 2003, 2004; Trujillo & Allen, 2007) during No-Go trials compared with Go trials.”

“Speci fi cally, the ERN is primarily due to increased theta-band (4 – 8 Hz) power (i.e., amplitude 2 ) and phase resetting (quanti fi ed as intertrial phase coherence [ITPC]) when making an error (42 – 44).”

“The link between theta and error processing is also supported by its relationship to the error-related negativity (ERN); an event-related potential (ERP) component that peaks after an erroneous response (Holroyd & Coles, 2002; Luu et al., 2004; Trujillo & Allen, 2007).”

“Theta oscillations exhibit maximal increases in power (magnitude) and synchrony (phase alignment) over MFC following error responses [18], [29], [31], [32].”

“As previous studies indicate increased synchronization associated with the ERN in the theta frequency band (4–7 Hz) and 25–75 ms time window [22, 24, 25, 26], all analysis was performed for this time and frequency range.”

“…et al., 1998; Pizzagalli, Peccoralo, Davidson, & Cohen, 2006; Rüsseler, Kuhlicke, & Münte, 2003), magnetoencephalography (Miltner et al., 2003), time-frequency analyses (Luu, Tucker, & Makeig, 2004; Trujillo & Allen, 2007), and intracerebral recording (Brázdil, Roman, Daniel, & Rektor, 2005).”

“Furthermore, cognitive neuroscience research has identified frontal midline theta as a potential neural mechanism involved in cognitive control processes, suggesting its role in facilitating top-down cognitive control across extensive brain networks [114], [115].”

“…in a broad range of cognitive tasks that require the allocation of attentional resources, and close relationship between event-related potentials (ERP), such as the error-related negativity (ERN) and evoked FMT oscillations has been reported (Trujillo and Allen 2007; van Driel et al. 2012).”