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.”

“…the medial frontal cortex (Cavanagh et al., 2009; Luu & Tucker, 2001; Luu, Tucker, Derryberry, Reed, & Poulsen, 2003; Luu, Tucker, & Makeig, 2004; Trujillo & Allen, 2007; Yordanova et al., 2004), as have the CRN (Burle, Roger, Allain, Vidal, & Hasbroucq, 2008; Cavanagh et al., 2009; Yordanova et…”

“Spectral decomposition has revealed that the ERN (errors), FRN (punishment), and N2 (conflict) share a common signature in the theta band (Cavanagh et al., 2009; Luu et al., 2004, 2003; Trujillo and Allen, 2007; Yordanova et al., 2004) (Fig.”

“Indeed, numerous studies have shown that FMT oscillations are also modulated by error-related signals (e.g., Luu et al., 2004; Trujillo and Allen, 2007) and reward expectation (Cohen et al., ), even though these tasks do not seem to have significant memory demands.”

“…and power enhancement over the medial frontal cortex (Bernat et al., 2008; Cavanagh et al., 2009; Cohen et al., 2007; Marco-Pallares et al., 2008; Trujillo and Allen, 2007), supporting the major postulate of Holroyd and Coles' (2002) reinforcement learning theory that these two ERPs reflect the…”

“5 are differences in other frequency bands implicated in cognitive control, such as theta (Cavanagh et al., 2009; Luu et al., 2004; Marco-Pallares et al., 2008; Trujillo and Allen, 2007).”

“Interestingly, it has been shown that erroneous manual responses elicit increases in both phase-and non-phase-locked theta activity (Trujillo and Allen, 2007).”

“…these aforementioned responselocked ERPs have also been shown to reflect phase-locked mid-frontal theta-band activities (Luu and Tucker 2001; Luu et al. 2004; Trujillo and Allen 2007), we are unaware of any investigations of the EEG correlates of response uncertainty during reinforcement learning.”

“…and conflict resolution (Cavanagh et al., 2009; Cohen, van Gaal, Ridderinkhof, & Lamme, 2009; Cohen, Ridderinkhof, Haupt, Elger, & Fell, 2008; Trujillo & Allen, 2007; Luu, Tucker, & Makeig, 2004), we aimed at test- ing whether processing of cognitive conflicts can be characterized according…”

“At the cognitive level, medial frontal theta has been linked to error monitoring and feedback processing during learning tasks (Bates et al., 2009; Cavanagh et al., 2009; Cohen et al., 2008b; Holroyd and Coles, 2002; Mazaheri et al., 2009; Trujillo and Allen, 2007; Wang et al., 2005).”

“…localization (Holroyd et al. 1998; Pizzagalli et al. 2006) magnetoencephalography (Miltner et al. 2003), time–frequency analyses (Luu et al. 2004; Trujillo and Allen, 2007), and intracerebral recording (Brázdil et al. 2005) suggest that the ERN is generated in the anterior cingulate cortex…”

“…activity in the theta (4 – 8 Hz) band may be the electrophysiological mechanism underlying these interactions (Luu and Tucker, 2001; Luu et al., 2004; Trujillo and Allen, 2007), reflected by midfrontal theta power and frontal intersite theta phase synchronization (Cavanagh et al., 2009).”

“…work has demonstrated that the error-related negativity (ERN; Gehring et al., 1993; Falkenstein et al., 1991) is well characterized by theta TF measures (Gehring and Willoughby, 2004; Trujillo and Allen, 2007; Yordanova et al., 2004; Cavanagh et al., 2009a, 2011; Cohen and Cavanagh, 2011).”

“…including the correct-and error-related negativity reflect a medial frontal theta oscillatory process that is both enhanced and partly phase-reset during these cognitive events [(Luu et al., 2004; Trujillo and Allen, 2007; Wang et al., 2005), but see (Yeung et al., 2007), for an alternative view].”

“…(Cohen, 2011; Cavanagh, Frank, Klein, & Allen, 2010; Cavanagh, Cohen, & Allen, 2009; Christie & Tata, 2009; Marco-Pallares et al., 2008, 2009; Cohen, Ridderinkhof, Haupt, Elger, & Fell, 2008; Cohen, Elger, & Ranganath, 2007; Trujillo & Allen, 2007; Luu, Tucker, & Makeig, 2004; Luu & Tucker, 2001).”

“Furthermore, increased theta synchronization has been associated with conflict-related processes, such as error detection and post-error correction (Cavanagh et al., 2009; Luu et al., 2004; Trujillo and Allen, 2007), goal conflict and response selection (Moore et al., 2006, 2012).”

“We thus used 2 automated procedures for assessing nonstationarity, KPSS and ADF, both from the Granger Causality Matlab toolbox (Seth 2010), which is available for free download on the internet.”

“, frontocentral theta and delta signals) following several different kinds of events, including action errors (Luu et al., 2004; Trujillo and Allen, 2007; Cohen, 2011; van Driel et al., 2012) and prediction errors (Cavanagh et al.”

“Recent work has drawn a link between FM theta and the error-related signals recorded in the evoked potential (Trujillo and Allen, 2007; Cohen et al., 2008; Womelsdorf et al., 2010a), or to prediction error signals arising in the context of reward-guided learning (Cavanagh et al.”

“…their own behaviors, including response conflicts (Cohen and Cavanagh, 2011; Cohen and Donner, 2013; Nigbur et al., 2011), error monitoring (Cavanagh et al., 2009; Luu et al., 2004; Trujillo and Allen, 2007), task switching (Cunillera et al., 2012), and feedback processing (Cavanagh et al., 2010).”

“While the results concerning enhancement of theta power is in keeping with most of the previous literature (Trujillo and Allen, 2007; Cavanagh et al., 2009; van Driel et al., 2012), the enhancement of alpha band in the very same conditions found for theta is less expected and deserves some…”

“Neural computations in the MFC, and its interactions with other cortical and subcortical areas, seem to be coordinated by electrophysiological oscillations in the theta (4–8 Hz) band [7,8,9,10,11,12,13,14,15].”

“The slow and punctate nature of the Rew-P and associated spectral activities suggest that this reward-related phenomenon may not be due to phase reset of an obligatory exogenously-invoked oscillation, such as has been postulated to underlie the N2/FRN and initial P3 at FCz (Luu et al., 2004; Trujillo and Allen, 2007;…”

“…previous EEG studies associated increases in DMPFC (4 – 8 Hz) power and DMPFC-DLPFC phase synchronization with conflict detection, response monitoring, and cognitive control (Trujillo and Allen, 2007; Cohen et al., 2008; Hanslmayr et al., 2008; Cavanagh et al., 2009, 2012; Womelsdorf et al., 2010).”

“…1997; ieuwenhuis, Slagter, et al., 2005; Ullsperger & von Cramon, 2003), erhaps due to partial phase resetting and amplitude enhancement f the frontal midline theta (Cavanagh, Cohen, & Allen, 2009; Luu, ucker, Derryberry, Reed, & Poulsen, 2003; Luu, Tucker, & Makeig, 004; Trujillo & Allen, 2007).”

“There is also growing evidence that theta oscillations are linked to error 572 monitoring (Cavanagh, Cohen, & Allen, 2009; Gevins & Smith, 2000; Luu, Tucker, & Makeig, 573 2004; Trujillo & Allen, 2007; Yordanova et al., 2004).”

“The ERN is thought to contain a combination of phase-locked theta-frequency-band activity and non-phase-locked theta-frequency-band power increases (Yeung et al., 2007; Trujillo and Allen, 2007; Wang et al., 2005; Luu et al., 2004).”

“As the ACC has previously been shown to be involved in error-detection and processing during cognitive tasks (Luu et al., 2004; Trujillo and Allen, 2007), it is suggested to subserve a similar process during balance control by detecting discrepancies between expected and actual state of balance…”

“…sites (typically maximal at electrode FCz), increases with conflict and errors, correlates with RT, and predicts post-error adjustments (Cavanagh et al., 2009; Cohen, 2011b; Cohen and Cavanagh, 2011; Hanslmayr et al., 2008; Nigbur et al., 2011; Trujillo and Allen, 2007; Yordanova et al., 2004).”

“The ERN is generated by theta band phase resetting and enhancement (Trujillo and Allen, 2007) in the ACC and surrounding mPFC, and is thought to reflect the functions of an action monitoring system that uses signals of error, conflict or punishment to adapt future behavior (Debener et al.”

“A limitation of our procedure is that it cannot distinguish between changes in the across-trial variability of ERP peak latencies versus changes in the rate at which ongoing prestimulus EEG oscillations are realigned following stimulation onset, a process known as phase-resetting (Gruber, Klimesch, Sauseng, and Doppel…”

“020 band (7,10) and can be interpreted as brief bursts or complexes of theta-band activity partially time- and phase-locked to relevant stimulus presentations (11) or motor responses (12).”

“While most research in frontoparietal oscillatory signatures of control focus on theta band dynamics (Cavanagh and Frank, 2014; Cooper et al., 2015a; Cunillera et al., 2012; Gladwin and de Jong, 2005; Luu et al., 2004; Moore et al., 2012; Sauseng et al., 2006; Trujillo and Allen, 2007), our findings provide novel insi…”

“For example, Luu et al. (2004) filtered the ERPs within a 4–12 Hz bandpass, while Trujillo and Allen (2007) used 3–13 Hz bandpass filter in order to optimize ERN component.”

“The topographical distributions of theta across all scalp-level spatial transformations are consistentwith many previous studies of error-related event-related potentials and theta-band activities (Cavanagh et al., 2012; Gehring et al., 2012; Luu et al., 2004; Trujillo and Allen, 2007).”

“…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).”

“In particular, whether the EEG signature of response conflict monitoring truly reflects a “neural oscillation” or simply a non-oscillatory transient remains debated (Yeung et al., 2007; Trujillo and Allen, 2007; Cohen and Donner, 2013).”

“, 2004) that is assumed to be determined by the dopaminergic system (Cavanagh et al., 2010; Trujillo and Allen, 2007).”

“Specifically, others have claimed that the theta response associated with errors could be the consequence of the mathematical transformation performed on the ERN during time-frequency analysis (Luu et al. 2004; Trujillo and Allen 2007).”

“, 1992; Demiralp and Basar, 1992; Karakas, 1997; Klimesch, 1999), error processing (Luu et al., 2003, 2004; Trujillo and Allen, 2007), reward processing (Cohen et al.”

“…Rangaswamy et al., 2007a), stroop effect (Hanslmayr et al., 2008), emotion processing (Krause et al., 2000b; Doppelmayr et al., 2002; Aftanas et al., 2003), error processing (e.g. Luu et al., 2004; Trujillo and Allen, 2007), and outcome processing (Gehring and Willoughby, 2004; Cohen et al., 2007).”

“Received 27 June 2011 Revised 3 August 2011 Accepted 11 August 2011 Available online 27 August 2011”

“Oscillatory and synchronization activities in three frequency ranges were analyzed (i) in the theta band (4–8 Hz), based upon its modulation of the ERN (Trujillo and Allen, 2007; Cavanagh et al., 2009; Luu et al., 2004); (ii) in the alpha band (8–13 Hz), specifically over occipital electrode regions, as an indicator o…”

“We were interested in power in the theta band as a measure of error monitoring (Luu et al., 2004; Debener et al., 2005; Trujillo and Allen, 2007), and power in the alpha band as a measure of post-error modulations in cortical arousal (Carp and Compton, 2009; Compton et al.”

“These findings are consistent with a large number of EEG studies suggesting that fm-theta increases in response to more difficult task requirements [6,7,12,15], response inhibition [3,4,5] and errors [8,46].”

“Studies show that ERN arises at least in part from ongoing theta activity of 4 – 7 Hz (Luu et al., 2003; Luu et al., 2004; Luu and Tucker, 2001; Trujillo and Allen, 2007).”

“Theta signatures in the EEG have been observed during error/conflict monitoring (Trujillo and Allen, 2007) and feedback processing (Cavanagh et al.”

“Several studies using intracerebral recordings in animals and humans concluded that fmtheta has its main source in the MCC (Wang et al., 2005; Tsujimoto et al., 2009; Womelsdorf et al., 2010).”

“…speaking, brain electrical oscillations occurring between 4 and 8 Hertz, called theta activity, have been associated with memory processing, such as during memory encoding [42], recognition or processing during spatial navigation [65] and other related processes including error detection [61].”

“…little functional importance of the phase-locked component of midfrontal theta (Cohen and Donner, 2013), although other studies have highlighted that the phase-locked component of error-related activitymay also provide some unique information (Munneke et al., 2015; Trujillo and Allen, 2007).”

“Furthermore, a greater extent of non-phase locked θ-activity was observed post-error-event in comparison to phase-locked activity [67],”

“…et al., 2007], Stroop effect [Hanslmayr et al., 2008], emotion processing [Aftanas et al., 2003; Doppelmayr et al., 2002; Krause et al., r 1030 r 2000b], error processing [e.g., Trujillo and Allen, 2007; Luu et al., 2004], and outcome processing [Cohen et al., 2007; Gehring and Willoughby, 2004].”

“For example, changes in theta-band and delta-band oscillations along the frontal midline coincide with the ERN 60–62 , with both phase resetting and changes of power in ongoing theta oscillations contributing to this signal 63,64 .”

“A prominent neurophysiological model suggests that the ACC, as part of the reinforcement learning network, gives rise to the FRN (Ridderinkhof, Ullsperger, Crone, & Nieuwenhuis, 2004) via a phaselocked enhancement of theta activity (Luu, Tucker, & Makeig, 2004; Trujillo & Allen, 2007).”

“Pre-filtering was done in order to directly target the relevant activity known to be associated with FRN and P3 (theta and delta, respectively; e.g., Bernat et al. 2007, 2011; Demiralp et al. 2001; Gehring and Willoughby 2004; Gilmore et al. 2010; Trujillo and Allen 2007) and because it yielded the most parsimonious T…”

“…Isomura et al., 2003; Womelsdorf et al., 2010), magnetoencephalography (MEG; Ishii et al., 1999), electroencephalography (EEG; Luu et al., 2004; Trujillo and Allen, 2007; van Noordt et al., 2015a), functional magnetic resonance imaging (fMRI; Meltzer et al., 2007), and transcranial direct…”

“(2009) examined scalp-recorded medial frontal theta activity (4-8 Hz; the frequency band thought to underlie the generation of the ERN; Luu et al., 2004; Trujillo and Allen, 2007) following the commission of errors.”