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The online version of the original article can be found under doi: 10.1007/s10459-015-9648-3 .

In recent years there has been an increase in the number of portable low-cost electroencephalographic (EEG) systems available to researchers. However, to date the validation of the use of low-cost EEG systems has focused on continuous recording of EEG data and/or the replication of large system EEG setups reliant on event-markers to afford examination of event-related brain potentials (ERP). Here, we demonstrate that it is possible to conduct ERP research without being reliant on event markers using a portable MUSE EEG system and a single computer. Specifically, we report the results of two experiments using data collected with the MUSE EEG system – one using the well-known visual oddball paradigm and the other using a standard reward-learning task. Our results demonstrate that we could observe and quantify the N200 and P300 ERP components in the visual oddball task and the reward positivity (the mirror opposite component to the feedback-related negativity) in the reward-learning task. Specifically, single sample t-tests of component existence (all p’s < 0.05), computation of Bayesian credible intervals, and 95% confidence intervals all statistically verified the existence of the N200, P300, and reward positivity in all analyses. We provide with this research paper an open source website with all the instructions, methods, and software to replicate our findings and to provide researchers with an easy way to use the MUSE EEG system for ERP research. Importantly, our work highlights that with a single computer and a portable EEG system such as the MUSE one can conduct ERP research with ease thus greatly extending the possible use of the ERP methodology to a variety of novel contexts.

Perceived ownership has been shown to impact a variety of cognitive processes: attention, memory, and—more recently—reward processing. In the present experiment we examined whether or not perceived ownership would interact with the construct of value—the relative worth of an object. Participants completed a simple gambling game in which they gambled either for themselves or for another while electroencephalographic data were recorded. In a key manipulation, gambles for oneself or for another were for either small or large rewards. We tested the hypothesis that value affects the neural response to self-gamble outcomes, but not other-gamble outcomes. Our experimental data revealed that while participants learned the correct response option for both self and other gambles, the reward positivity evoked by wins was impacted by value only when gambling for oneself. Importantly, our findings provide additional evidence for a self-ownership bias in cognitive processing and further demonstrate the insensitivity of the medial-frontal reward system to gambles for another.

Items that are produced (e.g., read aloud) during encoding typically are better remembered than items that are not produced (e.g., read silently). This "production effect" has been explained by distinctiveness: Produced items have more distinct features than nonproduced items, leading to enhanced retrieval. The goal of the current study was to use electroencephalography (EEG) to examine the neural basis of the production effect. During study, participants were presented with words that they were required to read silently, read aloud, or sing while EEG data were recorded. Subsequent memory performance was tested using a yes/no recognition test. Analysis focused on the event-related brain potentials (ERPs) evoked by the encoding instruction cue for each instruction condition. Our data revealed enhanced memory performance for produced items and a greater P300 ERP amplitude for instructions to sing or read aloud compared with instructions to read silently. Our results demonstrate that the amplitude of the P300 is modulated by at least 1 aspect of production, vocalization (singing/reading aloud relative to reading silently), and are consistent with the distinctiveness account of the production effect. The ERP methodology is a viable tool for investigating the production effect. (PsycINFO Database Record

Objects that are rare are often perceived to be inherently more valuable than objects that are abundant - a bias brought about in part by the scarcity heuristic. In the present study, we sought to test whether perception of rarity impacted reward evaluation within the human medial-frontal cortex. Here, participants played a gambling game in which they flipped rare and abundant 'cards' on a computer screen to win financial rewards while electroencephalographic data were recorded. Unbeknownst to participants, reward outcome and frequency was random and equivalent for both rare and abundant cards; thus, only a perception of scarcity was true. Analysis of the electroencephalographic data indicated that the P300 component of the event-related brain potential differed in amplitude for wins and losses following the selection of rare cards, but not following the selection of abundant cards. Importantly, then, we found that the perception of card rarity impacted reward processing even though reward feedback was independent of and subsequent to card selection. Our data indicate a top-down influence of the scarcity heuristic on reward evaluation, and specifically the processing of reward magnitude, within the human medial-frontal cortex.

We used the event-related potential (ERP) methodology to examine differences in neural processing between visually and memory-guided reaches. Consistent with previous findings (e.g., Westwood, Heath, & Roy, 2003), memory-guided reaches undershot veridical target location to a greater extent than their visually guided counterparts. Analysis of the ERP data revealed that memory-guided reaches were associated with reduced potentials over medial-frontal cortex at target presentation and following movement onset. Further, we found that the amplitudes of the potentials over medial-frontal cortex for visually and memory-guided reaches were significantly correlated with the peak accelerations and decelerations of the reaching movements. Our results suggest that memory-guided reaches are mediated by a motor plan that is generated while a target is visible, and then stored in memory until needed—a result counter to recent behavioral theories asserting that memory-guided reaches are planned just before movement onset via a stored, sensory-based target representation.

Neural correlates of cognitive states in event-related potentials (ERPs) serve as markers for related cerebral processes. Although these are usually evaluated in subject groups, the ability to evaluate such markers statistically in single subjects is essential for case studies in neuropsychology. Here we investigated the use of a simple test based on nonparametric bootstrap confidence intervals for this purpose, by evaluating three different ERP phenomena: the face-selectivity of the N170, error-related negativity, and the P3 component in a Posner cueing paradigm. In each case, we compare single-subject analysis with statistical significance determined using bootstrap to conventional group analysis using analysis of variance (ANOVA). We found that the proportion of subjects who show a significant effect at the individual level based on bootstrap varied, being greatest for the N170 and least for the P3. Furthermore, it correlated with significance at the group level. We conclude that the bootstrap methodology can be a viable option for interpreting single-case ERP amplitude effects in the right setting, probably with well-defined stereotyped peaks that show robust differences at the group level, which may be more characteristic of early sensory components than late cognitive effects.

The own-race bias (ORB) is a well-documented recognition advantage for own-race (OR) over cross-race (CR) faces, the origin of which remains unclear. In the current study, event-related potentials (ERPs) were recorded while Caucasian participants age-categorized Black and White faces which were digitally altered to display either a race congruent or incongruent facial structure. The results of a subsequent surprise memory test indicated that regardless of facial structure participants recognized White faces better than Black faces. Additional analyses revealed that temporally-early ERP components associated with face-specific perceptual processing (N170) and the individuation of facial exemplars (N250) were selectively sensitive to skin color. In addition, the N200 (a component that has been linked to increased attention and depth of encoding afforded to in-group and OR faces) was modulated by color and structure, and correlated with subsequent memory performance. However, the LPP component associated with the cognitive evaluation of perceptual input was influenced by racial differences in facial structure alone. These findings suggest that racial differences in skin color and facial structure are detected during the encoding of unfamiliar faces, and that the categorization of conspecifics as members of our social in-group on the basis of their skin color may be a determining factor in our ability to subsequently remember them.

There is disagreement about the effectiveness of observational practice for the acquisition of novel coordination skills and the type of processes involved in observation of novel movements. In this study, we examined learning of a bimanual 90 degrees phase offset through comparisons of three groups; physical practice, observational practice and no practice (n = 12/group). Groups were compared before and after practice on perception and production scans of the practised pattern. The observation group was yoked to the physical group such that observers watched repeated demonstrations of a learning model. Although there were no positive effects of observational practice for physical performance measures, the observation group did not differ from the physical practice group and was more accurate than controls on perceptual discrimination measures after practice. We concluded that observation of a novel bimanual movement can aid perception but that physical practice is necessary for immediate physical performance benefits. These results are discussed in terms of cognitive mediation models of motor skill learning.

A recent theory holds that the anterior cingulate cortex (ACC) uses reinforcement learning signals conveyed by the midbrain dopamine system to facilitate flexible action selection. According to this position, the impact of reward prediction error signals on ACC modulates the amplitude of a component of the event-related brain potential called the error-related negativity (ERN). The theory predicts that ERN amplitude is monotonically related to the expectedness of the event: It is larger for unexpected outcomes than for expected outcomes. However, a recent failure to confirm this prediction has called the theory into question. In the present article, we investigated this discrepancy in three trial-and-error learning experiments. All three experiments provided support for the theory, but the effect sizes were largest when an optimal response strategy could actually be learned. This observation suggests that ACC utilizes dopamine reward prediction error signals for adaptive decision making when the optimal behavior is, in fact, learnable.

The N200 and the feedback error-related negativity (fERN) are two components of the event-related brain potential (ERP) that share similar scalp distributions, time courses, morphologies, and functional dependencies, which raises the question as to whether they are actually the same phenomenon. To investigate this issue, we recorded the ERP from participants engaged in two tasks that independently elicited the N200 and fERN. Our results indicate that they are, in fact, the same ERP component and further suggest that positive feedback elicits a positive-going deflection in the time range of the fERN. Taken together, these results indicate that negative feedback elicits a common N200 and that modulation of fERN amplitude results from the superposition on correct trials of a positive-going deflection that we term the feedback correct-related positivity.

Different neural systems underlie the evaluation of different types of errors. Recent electroencephalographic evidence suggests that outcome errors -- errors indicating the failure to achieve a movement goal -- are evaluated within medial-frontal cortex (Krigolson and Holroyd 2006, 2007a, b). Conversely, evidence from a variety of manual aiming studies has demonstrated that target errors -- discrepancies between the actual and desired motor command brought about by an unexpected change in the movement environment -- are mediated within posterior parietal cortex (e.g., Desmurget et al. 1999, 2001; Diedrichsen et al. 2005). Here, event-related brain potentials (ERP) were recorded to assess medial-frontal and parietal ERP components associated with the evaluation of outcome and target errors during performance of a manual aiming task. In line with previous results (Krigolson and Holroyd 2007a), we found that target perturbations elicited an ERP component with a parietal scalp distribution, the P300. However, the timing of kinematic changes associated with accommodation of the target perturbations relative to the timing of the P300 suggests that the P300 component was not related to the online control of movement. Instead, we believe that the P300 evoked by target perturbations reflects the updating of an internal model of the movement environment. Our results also revealed that an error-related negativity, an ERP component typically associated with the evaluation of speeded response errors and error feedback, was elicited when participants missed the movement target. Importantly, this result suggests that a reinforcement learning system within medial-frontal cortex may play a role in improving subsequent motor output.

The authors manipulated the availability of monocular and binocular vision during the constituent planning and control stages of a goal-directed reaching task. Furthermore, trials were completed with or without online limb vision to determine whether monocular- or binocular-derived ego-motion cues influence the integration of visual feedback for online limb corrections. Results showed that the manipulation of visual cues during movement planning did not influence planning times or overall kinematics. During movement execution, however, binocular reaches—and particularly those completed with online limb vision—demonstrated heightened endpoint accuracy and stability, a finding directly linked to the adoption of a feedback-based mode of reaching control (i.e., online control). In contrast, reaches performed with online monocular vision produced increased endpoint error and instability and demonstrated reduced evidence of feedback-based corrections (i.e., offline control). Based on these results, the authors propose that the combination of static (i.e., target location) and dynamic (i.e., the moving limb) binocular cues serve to specifically optimize online reaching control. Moreover, results provide new evidence that differences in the kinematic and endpoint parameters of binocular and monocular reaches reflect differences in the extent to which the aforementioned engage in online and offline modes of movement control.

We have recently provided evidence that an error-related negativity (ERN), an ERP component generated within medial-frontal cortex, is elicited by errors made during the performance of a continuous tracking task (O.E. Krigolson & C.B. Holroyd, 2006). In the present study we conducted two experiments to investigate the ability of the medial-frontal error system to evaluate predictive error information. In two experiments participants used a joystick to perform a computer-based continuous tracking task in which some tracking errors were inevitable. In both experiments, half of these errors were preceded by a predictive cue. The results of both experiments indicated that an ERN-like waveform was elicited by tracking errors. Furthermore, in both experiments the predicted error waveforms had an earlier peak latency than the unpredicted error waveforms. These results demonstrate that the medial-frontal error system can evaluate predictive error information.

Self-referencing (i.e., thinking about oneself during encoding) can increase attention toward to-be-encoded material, and support memory for information in adults and children. The current inquiry tested an educational application of this ‘self reference effect’ (SRE) on memory. A self-referential modification of literacy tasks (vocabulary spelling) was tested in two experiments. In Experiment 1, seven-to nine-year-old children (N = 47) were asked to learn the spelling of four nonsense words by copying the vocabulary and generating sentences. Half of the children were asked to include themselves as a subject in each sentence. Results showed that children in this self-referent condition produced longer sentences and increased spelling accuracy by more than 20%, relative to those in an other-referent condition. Experiment 2 (N = 32) replicated this pattern in real-word learning. These findings demonstrate the significant potential advantages of utilizing self-referential encoding in the classroom.

Clinical reasoning is dependent upon working memory (WM). More precisely, during the clinical reasoning process stored information within long-term memory is brought into WM to facilitate the internal deliberation that affords a clinician the ability to reason through a case. In the present study, we examined the relationship between clinical reasoning and WM while participants read clinical cases with functional magnetic resonance imaging (fMRI). More specifically, we examined the impact of clinical case difficulty (easy, hard) and clinician level of expertise (2nd year medical students, senior gastroenterologists) on neural activity within regions of cortex associated with WM (i.e., the prefrontal cortex) during the reasoning process. fMRI was used to scan ten second-year medical students and ten practicing gastroenterologists while they reasoned through sixteen clinical cases [eight straight forward (easy) and eight complex (hard)] during a single 1-h scanning session. Within-group analyses contrasted the easy and hard cases which were then subsequently utilized for a between-group analysis to examine effects of expertise (novice > expert, expert > novice). Reading clinical cases evoked multiple neural activations in occipital, prefrontal, parietal, and temporal cortical regions in both groups. Importantly, increased activation in the prefrontal cortex in novices for both easy and hard clinical cases suggests novices utilize WM more so than experts during clinical reasoning. We found that clinician level of expertise elicited differential activation of regions of the human prefrontal cortex associated with WM during clinical reasoning. This suggests there is an important relationship between clinical reasoning and human WM. As such, we suggest future models of clinical reasoning take into account that the use of WM is not consistent throughout all clinical reasoning tasks, and that memory structure may be utilized differently based on level of expertise.

Perceptual judgments about the angular disparity of a character from its standard upright (i.e., mental rotation task) result in a concurrent increase in reaction time (RT) and modulation of the amplitude of the P300 event-related brain potential (ERP). It has therefore been proposed that the P300 represents the neural processes associated with a visual rotation. In turn, the visuomotor mental rotation (VMR) task requires reaching to a location that deviates from a target by a predetermined angle. Although the VMR task exhibits a linear increase in RT with increasing oblique angles of rotation, work has not examined whether the task is supported via a visual rotation analogous to its mental rotation task counterpart. This represents a notable issue because seminal work involving non-human primates has ascribed VMR performance to the motor-related rotation of directionally tuned neurons in the primary motor cortex. Here we examined the concurrent behavioral and ERP characteristics of a standard reaching task and VMR tasks of 35°, 70°, and 105° of rotation. Results showed that the P300 amplitude was larger for the standard compared to each VMR task--an effect independent of the angle of rotation. In turn, the amplitude of the contingent negative variation (CNV)--an ERP related to cognitive and visuomotor integration for movement preparation--was systematically modulated with angle of rotation. Thus, we propose that the CNV represents an ERP correlate related to the cognitive and/or visuomotor transformation demands of increasing the angular separation between a stimulus and a movement goal.

Prism adaptation (PA) is both a perceptual-motor learning task as well as a promising rehabilitation tool for visuo-spatial neglect (VSN) – a spatial attention disorder often experienced after stroke resulting in slowed and/or inaccurate motor responses to contralesional targets. During PA, individuals are exposed to prism-induced shifts of the visual-field while performing a visuo-guided reaching task. After adaptation, with goggles removed, visuo-motor responding is shifted to the opposite direction of that initially induced by the prisms. This visuo-motor aftereffect has been used to study visuo-motor learning and adaptation and has been applied clinically to reduce VSN severity by improving motor responding to stimuli in contralesional (usually left-sided) space. In order to optimize PA’s use for VSN patients, it is important to elucidate the neural and cognitive processes that alter visuomotor function during PA. In the present study, healthy young adults underwent PA while event-related potentials (ERPs) were recorded at the termination of each reach (screen-touch), then binned according to accuracy (hit vs. miss) and phase of exposure block (early, middle, late). Results show that two ERP components were evoked by screen-touch: an early error-related negativity (ERN), and a P300. The ERN was consistently evoked on miss trials during adaptation, while the P300 amplitude was largest during the early phase of adaptation for both hit and miss trials. This study provides evidence of two neural signals sensitive to visual feedback during PA that may sub-serve changes in visuomotor responding. Prior ERP research suggests that the ERN reflects an error processing system in medial-frontal cortex, while the P300 is suggested to reflect a system for context updating and learning. Future research is needed to elucidate the role of these ERP components in improving visuomotor responses among individuals with VSN.

The neural systems that afford our ability to evaluate rewards and punishments are impacted by a variety of external factors. Here, we demonstrate that increased cognitive load reduces the functional efficacy of a reward processing system within the human medial-frontal cortex. In our paradigm, two groups of participants used performance feedback to estimate the exact duration of one second while electroencephalographic (EEG) data was recorded. Prior to performing the time estimation task, both groups were instructed to keep their eyes still and avoid blinking in line with well established EEG protocol. However, during performance of the time-estimation task, one of the two groups was provided with trial-to-trial-feedback about their performance on the time-estimation task and their eye movements to induce a higher level of cognitive load relative to participants in the other group who were solely provided with feedback about the accuracy of their temporal estimates. In line with previous work, we found that the higher level of cognitive load reduced the amplitude of the feedback-related negativity, a component of the human event-related brain potential associated with reward evaluation within the medial-frontal cortex. Importantly, our results provide further support that increased cognitive load reduces the functional efficacy of a neural system associated with reward processing.

A brief, group cognitive behavioural therapy with running as the interoceptive exposure (IE; exposure to physiological sensations) component was effective in decreasing anxiety sensitivity (AS; fear of arousal sensations) levels in female undergraduates (Watt et al., Anxiety and Substance Use Disorders: The Vicious Cycle of Comorbidity, 201-219, 2008). Additionally, repeated exposure to running resulted in decreases in cognitive (i.e., catastrophic thoughts) and affective (i.e., feelings of anxiety) reactions to running over time for high AS, but not low AS, participants (Sabourin et al., "Physical exercise as interoceptive exposure within a brief cognitive-behavioral treatment for anxiety-sensitive women", Journal of Cognitive Psychotherapy, 22:302-320, 2008). A follow-up study including the above-mentioned intervention with an expanded IE component also resulted in decreases in AS levels (Sabourin et al., under review). The goals of the present process study were (1) to replicate the original process study, with the expanded IE component, and (2) to determine whether decreases in cognitive, affective, and/or somatic (physiological sensations) reactions to running would be related to decreases in AS. Eighteen high AS and 10 low AS participants completed 20 IE running trials following the 3-day group intervention. As predicted, high AS participants, but not low AS participants, experienced decreases in cognitive, affective, and somatic reactions to running over time. Furthermore, decreases in cognitive and affective, but not in somatic, reactions to running were related to decreases in AS levels. These results suggest that the therapeutic effects of repeated exposure to running in decreasing sensitivity to anxiety-related sensations are not related to decreasing the experience of somatic sensations themselves. Rather, they are related to altering the cognitive and affective reactions to these sensations.

In advance of grasping a visual object embedded within fins-in and fins-out Müller-Lyer (ML) configurations, participants formulated a premovement grip aperture (GA) based on the size of a neutral preview object. Preview objects were smaller, veridical, or larger than the size of the to-be-grasped target object. As a result, premovement GA associated with the small and large preview objects required significant online reorganization to appropriately grasp the target object. We reasoned that such a manipulation would provide an opportunity to examine the extent to which the visuomotor system engages egocentric and/or allocentric visual cues for the online, feedback-based control of action. It was found that the online reorganization of GA was reliably influenced by the ML figures (i.e., from 20 to 80% of movement time), regardless of the size of the preview object, albeit the small and large preview objects elicited more robust illusory effects than the veridical preview object. These results counter the view that online grasping control is mediated by absolute visual information computed with respect to the observer (e.g., Glover in Behav Brain Sci 27:3-78, 2004; Milner and Goodale in The visual brain in action 1995). Instead, the impact of the ML figures suggests a level of interaction between egocentric and allocentric visual cues in online action control.

Motor error evaluation appears to be a hierarchically organized process subserved by 2 distinct systems: a higher level system within medial-frontal cortex responsible for movement outcome evaluation (high-level error evaluation) and a lower level posterior system(s) responsible for the mediation of within-movement errors (low-level error evaluation). While a growing body of evidence suggests that a reinforcement learning system within medial-frontal cortex plays a crucial role in the evaluation of high-level errors made during discrete reaching movements and continuous motor tracking, the role of this system in postural control is currently unclear. Participants learned a postural control task via a feedback-driven trial-and-error shaping process. In line with previous findings, electroencephalographic recordings revealed that feedback about movement outcomes elicited a feedback error-related negativity: a component of the human event-related brain potential associated with high-level outcome evaluation within medial-frontal cortex. Thus, the data provide evidence that a high-level error-evaluation system within medial-frontal cortex plays a key role in learning to control our body posture.

The execution of an antisaccade selectively increases the reaction time (RT) of a subsequent prosaccade (the unidirectional prosaccade switch-cost). To explain this finding, the task-set inertia hypothesis asserts that an antisaccade requires a cognitively mediated non-standard task-set that persists inertially and delays the planning of a subsequent prosaccade. The present study sought to directly test the theoretical tenets of the task-set inertia hypothesis by examining the concurrent behavioural and the event-related brain potential (ERP) data associated with the unidirectional prosaccade switch-cost. Participants pseudo-randomly alternated between pro- and antisaccades while electroencephalography (EEG) data were recorded. As expected, the completion of an antisaccade selectively increased the RT of a subsequent prosaccade, whereas the converse switch did not influence RTs. Thus, the behavioural results demonstrated the unidirectional prosaccade switch-cost. In terms of the ERP findings, we observed a reliable change in the amplitude of the P3 - time-locked to task-instructions - when trials were switched from a prosaccade to an antisaccade; however, no reliable change was observed when switching from an antisaccade to a prosaccade. This is a salient finding because extensive work has shown that the P3 provides a neural index of the task-set required to execute a to-be-completed response. As such, results showing that prosaccades completed after antisaccades exhibited increased RTs in combination with a P3 amplitude comparable to antisaccades provides convergent evidence that the unidirectional prosaccade switch-cost is attributed to the persistent activation of a non-standard antisaccade task-set.

Biological systems are capable of learning that certain stimuli are valuable while ignoring the many that are not, and thus perform feature selection. In machine learning, one effective feature selection approach is the least absolute shrinkage and selection operator (LASSO) form of regularization, which is equivalent to assuming a Laplacian prior distribution on the parameters. We review how such Bayesian priors can be implemented in gradient descent as a form of weight decay, which is a biologically plausible mechanism for Bayesian feature selection. In particular, we describe a new prior that offsets or "raises" the Laplacian prior distribution. We evaluate this alongside the Gaussian and Cauchy priors in gradient descent using a generic regression task where there are few relevant and many irrelevant features. We find that raising the Laplacian leads to less prediction error because it is a better model of the underlying distribution. We also consider two biologically relevant online learning tasks, one synthetic and one modeled after the perceptual expertise task of Krigolson et al. (2009). Here, raising the Laplacian prior avoids the fast erosion of relevant parameters over the period following training because it only allows small weights to decay. This better matches the limited loss of association seen between days in the human data of the perceptual expertise task. Raising the Laplacian prior thus results in a biologically plausible form of Bayesian feature selection that is effective in biologically relevant contexts.

Our ability to make decisions is predicated upon our knowledge of the outcomes of the actions available to us. Reinforcement learning theory posits that actions followed by a reward or punishment acquire value through the computation of prediction errors—discrepancies between the predicted and the actual reward. A multitude of neuroimaging studies have demonstrated that rewards and punishments evoke neural responses that appear to reflect reinforcement learning prediction errors [e.g., Krigolson, O. E., Pierce, L. J., Holroyd, C. B., & Tanaka, J. W. Learning to become an expert: Reinforcement learning and the acquisition of perceptual expertise. Journal of Cognitive Neuroscience, 21, 1833–1840, 2009; Bayer, H. M., & Glimcher, P. W. Midbrain dopamine neurons encode a quantitative reward prediction error signal. Neuron, 47, 129–141, 2005; O'Doherty, J. P. Reward representations and reward-related learning in the human brain: Insights from neuroimaging. Current Opinion in Neurobiology, 14, 769–776, 2004; Holroyd, C. B., & Coles, M. G. H. The neural basis of human error processing: Reinforcement learning, dopamine, and the error-related negativity. Psychological Review, 109, 679–709, 2002]. Here, we used the brain ERP technique to demonstrate that not only do rewards elicit a neural response akin to a prediction error but also that this signal rapidly diminished and propagated to the time of choice presentation with learning. Specifically, in a simple, learnable gambling task, we show that novel rewards elicited a feedback error-related negativity that rapidly decreased in amplitude with learning. Furthermore, we demonstrate the existence of a reward positivity at choice presentation, a previously unreported ERP component that has a similar timing and topography as the feedback error-related negativity that increased in amplitude with learning. The pattern of results we observed mirrored the output of a computational model that we implemented to compute reward prediction errors and the changes in amplitude of these prediction errors at the time of choice presentation and reward delivery. Our results provide further support that the computations that underlie human learning and decision-making follow reinforcement learning principles.

Ownership is a powerful construct. Indeed, in a series of recent studies, perceived ownership has been shown to increase attentional capacity, facilitate a memorial advantage, and elicit positive attitudes. Here, we sought to determine whether self-relevance would bias reward evaluation systems within the brain. To accomplish this, we had participants complete a simple gambling task during which they could “win” or “lose” prizes for themselves or for someone else, while electroencephalographic data were recorded. Our results indicated that the amplitude of the feedback error-related negativity, a component of the event-related brain potential sensitive to reward evaluation, was diminished when participants were not gambling for themselves. Furthermore, our data suggest that the ownership cues that indicated who would win or lose a given gamble either were processed as a potential for an increase in utility (i.e., gain: self-gambles) or were processed in a nonutilitarian manner (other-gambles). Importantly, our results suggest that the medial-frontal reward system is sensitive to perceived ownership, to the extent that it may not process changes in utility when they are not directly relevant to self.

Mind wandering episodes have been construed as periods of "stimulus-independent" thought, where our minds are decoupled from the external sensory environment. In two experiments, we used behavioral and event-related potential (ERP) measures to determine whether mind wandering episodes can also be considered as periods of "response-independent" thought, with our minds disengaged from adjusting our behavioral outputs. In the first experiment, participants performed a motor tracking task and were occasionally prompted to report whether their attention was "on-task" or "mind wandering." We found greater tracking error in periods prior to mind wandering vs. on-task reports. To ascertain whether this finding was due to attenuation in visual perception per se vs. a disruptive effect of mind wandering on performance monitoring, we conducted a second experiment in which participants completed a time-estimation task. They were given feedback on the accuracy of their estimations while we recorded their EEG, and were also occasionally asked to report their attention state. We found that the sensitivity of behavior and the P3 ERP component to feedback signals were significantly reduced just prior to mind wandering vs. on-task attentional reports. Moreover, these effects co-occurred with decreases in the error-related negativity elicited by feedback signals (fERN), a direct measure of behavioral feedback assessment in cortex. Our findings suggest that the functional consequences of mind wandering are not limited to just the processing of incoming stimulation per se, but extend as well to the control and adjustment of behavior.

Mirror-symmetrical reaching movements (i.e., antipointing) produce a visual-field-specific pattern of endpoint bias consistent with a perceptual representation of visual space (Heath et al. in Exp Brain Res 192:275–286, 2009a; J Mot Behav 41:383–392 2009b). The goal of the present investigation was to examine the concurrent behavioural and event-related brain potentials (ERP) of pro- and antipointing to determine whether endpoint bias in the latter task is related to a remapping of the environmental parameters of a target (i.e., vector inversion hypothesis) or a shift of visual attention from a veridical to a cognitively represented target location (i.e., reallocation of attention hypothesis). As expected, results for antipointing—but not propointing—yielded a visual-field-specific pattern of endpoint bias. In terms of the ERP findings, an early component (i.e., the N100) related to the orienting of visuospatial attention was comparable across pro- and antipointing. In contrast, a later occurring component (i.e., the P300) demonstrated a reliable between-task difference in amplitude. Notably, the P300 has been linked to the revision of a ‘mental model’ when a mismatch is noted between a stimulus and a required task goal (so-called context-updating). Thus, we propose that the between-task difference in the P300 indicates that antipointing is associated with a remapping of a target’s veridical location in mirror-symmetrical space (i.e., vector inversion). Moreover, our combined behavioural and ERP findings provide evidence that vector inversion is mediated via perception-based visual networks.

Previous research has demonstrated that higher-order cognitive processes associated with the allocation of selective attention are engaged when highly familiar self-relevant items are encountered, such as one's name, face, personal possessions and the like. The goal of our study was to determine whether these effects on attentional processing are triggered on-line at the moment self-relevance is established. In a pair of experiments, we recorded ERPs as participants viewed common objects (e.g., apple, socks, and ketchup) in the context of an “ownership” paradigm, where the presentation of each object was followed by a cue indicating whether the object nominally belonged either to the participant (a “self” cue) or the experimenter (an “other” cue). In Experiment 1, we found that “self” ownership cues were associated with increased attentional processing, as measured via the P300 component. In Experiment 2, we replicated this effect while demonstrating that at a visual–perceptual level, spatial attention became more narrowly focused on objects owned by self, as measured via the lateral occipital P1 ERP component. Taken together, our findings indicate that self-relevant attention effects are triggered by the act of taking ownership of objects associated with both perceptual and postperceptual processing in cortex.

To elucidate the neural mechanisms underlying the development of perceptual expertise, we recorded ERPs while participants performed a categorization task. We found that as participants learned to discriminate computer generated “blob” stimuli, feedback modulated the amplitude of the error-related negativity (ERN)—an ERP component thought to reflect error evaluation within medial–frontal cortex. As participants improved at the categorization task, we also observed an increase in amplitude of an ERP component associated with object recognition (the N250). The increase in N250 amplitude preceded an increase in amplitude of an ERN component associated with internal error evaluation (the response ERN). Importantly, these electroencephalographic changes were not observed for participants who failed to improve on the categorization task. Our results suggest that the acquisition of perceptual expertise relies on interactions between the posterior perceptual system and the reinforcement learning system involving medial–frontal cortex.

In the present experiment we investigated the impact of cognitive load on feedback evaluation by a learning system within medial-frontal cortex. Participants completed a task in which they had to use feedback to learn to accurately estimate the duration of one second. In two experimental conditions, we manipulated the cognitive load of the feedback stimuli (low load versus high load). Our results revealed that the amplitude of the feedback error-related negatvity (fERN), a component of the event-related brain potential (ERP) thought to index a learning system within medial-frontal, was reduced in the high load condition. Further, an analysis of the behavioural data revealed that in the high load condition participants made less effective adjustments to their estimates following error feedback. Taken together, our data suggest that the functional efficacy of the medial-frontal learning system is reduced as the cognitive load of feeback signals increase. Moreover, our data indicate that the effect of increased cognitive load is to increase the trial-to-trial temporal variability of feedback stimulus evaluation.

A recent theory holds that a component of the human event-related brain potential called the reward positivity reflects a reward prediction error signal. We investigated this idea in gambling-like task in which, on each trial, a visual stimulus predicted a subsequent rewarding or nonrewarding outcome with 80% probability. Consistent with earlier results, we found that the reward positivity was larger to unexpected than to expected outcomes. In addition, we found that the predictive cues also elicited a reward positivity, as proposed by the theory. These results indicate that the reward positivity reflects the initial assessment of whether a trial will end in success or failure and the reappraisal of that information once the outcome actually occurs.

Studies investigating the specificity hypothesis have not always demonstrated that reliance on a specific source of feedback increases with practice. The goal of the present study was to address this inconsistency by having participants practice a throwing task with or without vision at incremental levels (10, 50, 100, or 200 acquisition trials). Following acquisition, all participants in the present experiment performed 10 trials in a no-vision transfer condition. Our results demonstrated that, given a sufficient number of acquisition trials, feedback reliance increased as a function of time engaged in practice. Our results also suggest that increased reliance on a specific source of feedback occurs only after the control strategy for a task is optimized.

The feedback error-related negativity (fERN) is a component of the human event-related brain potential (ERP) elicited by feedback stimuli. A recent theory holds that the fERN indexes a reward prediction error signal associated with the adaptive modification of behavior. Here we present behavioral and ERP data recorded from participants engaged in a modified time estimation task. As predicted by the theory, our results indicate that fERN amplitude reflects a reward prediction error signal and that the size of this error signal is correlated across participants with changes in task performance.

The control of goal-directed reaching movements is thought to rely upon egocentric visual information derived from the visuomotor networks of the dorsal visual pathway. However, recent research (Krigolson and Heath, 2004) suggests it is also possible to make allocentric comparisons between a visual background and a target object to facilitate reaching accuracy. Here we sought to determine if the effectiveness of these allocentric comparisons is reduced as distance between a visual background and a target object increases. To accomplish this, participants completed memory-guided reaching movements to targets presented in an otherwise empty visual background or positioned within a proximal, medial, or distal visual background. Our results indicated that the availability of a proximal or medial visual background reduced endpoint variability relative to reaches made without a visual background. Interestingly, we found that endpoint variability was not reduced when participants reached to targets framed within a distal visual background. Such findings suggest that allocentric visual information is used to facilitate reaching performance; however, the fidelity by which such cues are used appears linked to the proximity of veridical target location. Importantly, these data also suggest that information from both the dorsal and ventral visual streams can be integrated to facilitate the online control of reaching movements.

Error processing during motor control involves the evaluation of “high-level” errors (i.e., failures to meet a system goal) by a frontal system involving anterior cingulate cortex and the evaluation of “low-level” errors (i.e., discrepancies between actual and desired motor commands) by a posterior system involving posterior parietal cortex. We have recently demonstrated that high-level errors committed within the context of a continuous tracking task elicited an error-related negativity (ERN) – a component of the event-related brain potential (ERP) generated within medial–frontal cortex that is sensitive to error commission. The purpose of the present study was to demonstrate that low-level motor errors do not elicit an ERN, but may instead evoke other ERP components associated with visual processing and online motor control. Participants performed a computer aiming task in which they manipulated a joystick to move a cursor from a start to a target position. On a random subset of trials the target jumped to a new position at movement onset, requiring the participants to modify their current motor command. Further, on one half of these “target perturbation” trials the cursor did not respond to corrective movements of the joystick. Consistent with our previous findings, we found that the uncorrectable errors elicited an ERN. We also found that the target perturbations on both correctable and uncorrectable trials did not elicit an ERN, but rather evoked two other ERP components, the N100 and P300. These results suggest that medial–frontal cortex is insensitive to low-level motor errors, and are in line with a recent theory that holds that the P300 reflects stimulus–response optimization by the impact of locus coeruleus activity on posterior cortex.

It has been proposed that visually guided reaching movements performed in the lower visual field (LVF) of peripersonal space are more effective and efficient than their upper visual field (UVF) counterparts (Danckert and Goodale 2001). In the present investigation we sought to determine whether this purported visual field asymmetry reflects advantaged processing of online visual feedback. To accomplish that objective, participants performed discrete reaching movements to each of three target locations in the LVF and UVF. In addition, reaches were completed under conditions wherein target location remained constant throughout a reaching response (i.e., control trials) and a separate condition wherein target location unexpectedly perturbed at movement onset (i.e., experimental trials). We reasoned that the target perturbation paradigm would provide a novel means to assess a possible superior–inferior visual field asymmetry for online reaching control. In terms of the impact of a target perturbation, both visual fields demonstrated equal proficiency integrating visual feedback for online limb adjustments. Interestingly, however, the spatial distribution of movement endpoints in the LVF was less than UVF counterparts (cf. Binsted and Heath 2005). Taken together, the present findings suggest that although LVF and UVF reaches readily use visual feedback to accommodate an unexpected target perturbation, reaches in the LVF elicit advantaged spatial benefits influencing the effectiveness of online limb corrections.

Human goal-directed behavior depends on multiple neural systems that monitor and correct for different types of errors. For example, tracking errors in continuous motor tasks appear to be processed by a system involving posterior parietal cortex, whereas errors in speeded response and trial-and-error learning tasks appear to be processed by a system involving frontal-medial cortex. To date, it is unknown whether there is a functional relationship between the posterior and frontal error systems. We recorded the event-related brain potential from participants engaged in a tracking task to investigate the role of the frontal system in continuous motor control. Our results demonstrate that tracking errors elicit temporally distinct error-related event-related brain potentials over frontal and posterior regions of the scalp, suggesting an interaction between the subcomponents of a hierarchically organized system for error processing. Specifically, we propose that the frontal error system assesses high-level errors (i.e. goal attainment) whereas the posterior error system is responsible for evaluating low-level errors (i.e. trajectory deviations during motor control).

Recent research [e.g., Carrozzo, M., Stratta, F., McIntyre, J., & Lacquaniti, F. (2002). Cognitive allocentric representations of visual space shape pointing errors. Experimental Brain Research 147 , 426–436; Lemay, M., Bertrand, C. P., & Stelmach, G. E. (2004). Pointing to an allocentric and egocentric remembered target. Motor Control , 8 , 16–32] reported that egocentric and allocentric visual frames of reference can be integrated to facilitate the accuracy of goal-directed reaching movements. In the present investigation, we sought to specifically examine whether or not a visual background can facilitate the online, feedback-based control of visually-guided (VG), open-loop (OL), and memory-guided (i.e. 0 and 1000 ms of delay: D0 and D1000) reaches. Two background conditions were examined in this investigation. In the first background condition, four illuminated LEDs positioned in a square surrounding the target location provided a context for allocentric comparisons (visual background: VB). In the second condition, the target object was singularly presented against an empty visual field (no visual background: NVB). Participants ( N = 14) completed reaching movements to three midline targets in each background (VB, NVB) and visual condition (VG, OL, D0, D1000) for a total of 240 trials. VB reaches were more accurate and less variable than NVB reaches in each visual condition. Moreover, VB reaches elicited longer movement times and spent a greater proportion of the reaching trajectory in the deceleration phase of the movement. Supporting the benefit of a VB for online control, the proportion of endpoint variability explained by the spatial location of the limb at peak deceleration was less for VB as opposed to NVB reaches. These findings suggest that participants are able to make allocentric comparisons between a VB and target (visible or remembered) in addition to egocentric limb and VB comparisons to facilitate online reaching control.