Publications by authors named "Olave E Krigolson"

39 Publications

Neural correlates of the production effect: An fMRI study.

Brain Cogn 2021 Jun 12;152:105757. Epub 2021 Jun 12.

Memorial University of Newfoundland, Department of Psychology, St. John's, NL A1B 3X9, Canada. Electronic address:

Recognition memory is improved for items produced at study (e.g., by reading them aloud) relative to a non-produced control condition (e.g., silent reading). This production effect is typically attributed to the extra elements in the production task (e.g., motor activation, auditory perception) enhancing item distinctiveness. To evaluate this claim, the present study examined the neural mechanisms underlying the production effect. Prior to a recognition memory test, different words within a study list were read either aloud, silently, or while saying "check" (as a sensorimotor control condition). Production improved recognition, and aloud words yielded higher rates of both recollection and familiarity judgments than either silent or control words. During encoding, fMRI revealed stronger activation in regions associated with motor, somatosensory, and auditory processing for aloud items than for either silent or control items. These activations were predictive of recollective success for aloud items at test. Together, our findings are compatible with a distinctiveness-based account of the production effect, while also pointing to the possible role of other processing differences during the aloud trials as compared to silent and control.
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http://dx.doi.org/10.1016/j.bandc.2021.105757DOI Listing
June 2021

Using EEG to decode semantics during an artificial language learning task.

Brain Behav 2021 Jun 15. Epub 2021 Jun 15.

Departments of Computing Science and Psychology, University of Alberta, Edmonton, Canada.

Background: As we learn a new nonnative language (L2), we begin to build a new map of concepts onto orthographic representations. Eventually, L2 can conjure as rich a semantic representation as our native language (L1). However, the neural processes for mapping a new orthographic representation to a familiar meaning are not well understood or characterized.

Methods: Using electroencephalography and an artificial language that maps symbols to English words, we show that it is possible to use machine learning models to detect a newly formed semantic mapping as it is acquired.

Results: Through a trial-by-trial analysis, we show that we can detect when a new semantic mapping is formed. Our results show that, like word meaning representations evoked by a L1, the localization of the newly formed neural representations is highly distributed, but the representation may emerge more slowly after the onset of the symbol. Furthermore, our mapping of word meanings to symbols removes the confound of the semantics to the visual characteristics of the stimulus, a confound that has been difficult to disentangle previously.

Conclusion: We have shown that the L1 semantic representation conjured by a newly acquired L2 word can be detected using decoding techniques, and we give the first characterization of the emergence of that mapping. Our work opens up new possibilities for the study of semantic representations during L2 learning.
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http://dx.doi.org/10.1002/brb3.2234DOI Listing
June 2021

The role of cognitive control and top-down processes in object affordances.

Atten Percept Psychophys 2021 Jul 26;83(5):2017-2032. Epub 2021 Mar 26.

Centre for Biomedical Research, University of Victoria, P.O. Box 1700 STN CSC, Victoria, British Columbia, V8W 2Y2, Canada.

A widely held though debatable claim is that the picture of an object like a frying pan automatically elicits features of a left/right-handed grasp action even in perceptual tasks that make no demands on the observer to consider the graspable properties of the depicted object. Here, we sought to further elucidate this claim by relying on a methodology that allowed us to distinguish between the influence of motor versus spatial codes on the selection of a left/right-handed response while electroencephalographic data were recorded. In our experiment, participants classified images of frying pans as upright or inverted using a left/right key press or by making a left/right-handed reach-and-grasp action towards a centrally located response element while we recorded electroencephalographic (EEG) data. In line with previous evidence (Bub, Masson, & van Noordenne, Journal of Experiment Psychology: Human Perception and Performance, 47(1), 53-80, 2021), these two modes of responding generated distinct correspondence effects on performance induced by the same set of images. In terms of our EEG data, we found that neither motor (the lateralized readiness potential) nor visual (N100 and P100) potentials were sensitive to handle-response hand correspondence. However, an exploratory theta analysis revealed that changes in frontal theta power mirrored the different correspondence effects evoked by the image on key press responses versus reach and grasp actions. Importantly, our results provide a link between these disparate effects and the engagement of cognitive control, highlighting a possible role of top-down control processes in separating motor features from the task-irrelevant features of an object, and thus in claims regarding object affordances more generally.
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http://dx.doi.org/10.3758/s13414-021-02296-zDOI Listing
July 2021

What happens when right means wrong? The impact of conflict arising from competing feedback responses.

Brain Res 2021 Feb 25;1761:147393. Epub 2021 Feb 25.

Theoretical and Applied Neuroscience Laboratory, University of Victoria, Canada.

Humans often rely on feedback to learn. Indeed, in learning the difference between feedback and an expected outcome is computed to inform future actions. Further, recent work has found that reward and feedback have a unique role in modulating conflict processing and cognitive control. However, it is still not clear how conflict, especially concerning the processing and evaluation of feedback, impacts learning. To address this, we examined the effects of feedback competition on feedback evaluation in a reinforcement learning task. Specifically, we had participants play a simple two-choice gambling game while electroencephalographic (EEG) data were recorded. On half of the experiment blocks, we reversed the meaning of performance feedback for each trial from its prepotent meaning to induce response conflict akin to the Stroop effect (e.g., '✓' meant incorrect). Behaviourally, we found that participants' accuracy was reduced as a result of incongruent feedback. Paralleling this, an analysis of our EEG revealed that incongruent feedback resulted in a reduction in amplitude of the reward positivity and the P300, components of the human event-related brain potential implicated in reward processing. Our results demonstrate the negative impact of conflict on feedback evaluation and the impact of this on subsequent performance.
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http://dx.doi.org/10.1016/j.brainres.2021.147393DOI Listing
February 2021

Dissociated neural signals of conflict and surprise in effortful decision Making: Theta activity reflects surprise while alpha and beta activity reflect conflict.

Neuropsychologia 2021 05 18;155:107793. Epub 2021 Feb 18.

Centre for Biomedical Research, University of Victoria, Victoria, British Columbia, V8W 2Y2, Canada.

What makes a decision difficult? Two key factors are conflict and surprise: conflict emerges with multiple competing responses and surprise occurs with unexpected events. Conflict and surprise, however, are often thought of as parsimonious accounts of decision making rather than an integrated narrative. We sought to determine whether conflict and/or surprise concurrently or independently elicit effortful decision making. Participants made a series of diagnostic decisions from physiological readings while electroencephalographic (EEG) data were recorded. To induce conflict and surprise, we manipulated task difficulty by varying the distance between a presented physiological reading and the category border that separated the two diagnoses. Whereas frontal theta oscillations reflected surprise - when presented readings were far from the expected mean, parietal alpha and beta oscillations indicated conflict - when readings were near the category border. Our findings provide neural evidence that both conflict and surprise engage cognitive control to employ effort in decision making.
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http://dx.doi.org/10.1016/j.neuropsychologia.2021.107793DOI Listing
May 2021

Using Muse: Rapid Mobile Assessment of Brain Performance.

Front Neurosci 2021 28;15:634147. Epub 2021 Jan 28.

Faculty of Health and Social Development, University of British Columbia Okanagan, Kelowna, BC, Canada.

The advent of mobile electroencephalography (mEEG) has created a means for large scale collection of neural data thus affording a deeper insight into cognitive phenomena such as cognitive fatigue. Cognitive fatigue - a neural state that is associated with an increased incidence of errorful performance - is responsible for accidents on a daily basis which at times can cost human lives. To gain better insight into the neural signature of cognitive fatigue in the present study we used mEEG to examine the relationship between perceived cognitive fatigue and human-event related brain potentials (ERPs) and electroencephalographic (EEG) oscillations in a sample of 1,000 people. As a secondary goal, we wanted to further demonstrate the capability of mEEG to accurately measure ERP and EEG data. To accomplish these goals, participants performed a standard visual oddball task on an Apple iPad while EEG data were recorded from a Muse EEG headband. Counter to traditional EEG studies, experimental setup and data collection was completed in less than seven minutes on average. An analysis of our EEG data revealed robust N200 and P300 ERP components and neural oscillations in the delta, theta, alpha, and beta bands. In line with previous findings we observed correlations between ERP components and EEG power and perceived cognitive fatigue. Further, we demonstrate here that a linear combination of ERP and EEG features is a significantly better predictor of perceived cognitive fatigue than any ERP or EEG feature on its own. In sum, our results provide validation of mEEG as a viable tool for research and provide further insight into the impact of cognitive fatigue on the human brain.
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http://dx.doi.org/10.3389/fnins.2021.634147DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7876403PMC
January 2021

Distribution of Practice Combined with Observational Learning Has Time Dependent Effects on Motor Skill Acquisition.

Percept Mot Skills 2021 Apr 17;128(2):885-899. Epub 2020 Dec 17.

Faculty of Health and Social Development, University of British Columbia Okanagan, Kelowna, Canada.

Studies of the benefits of a distributed practice schedule on motor skill acquisition have typically found that distribution of practice results in better learning. However, less research has focused on how the benefits of distributed practice are impacted by timing during acquisition. To examine how timing of skill acquisition interacts with distribution of practice we had two groups of participants complete either an extensive massed or distributed training schedule to learn a speed stacking sequence across ten sessions. For participants in both groups, we provided observational learning to facilitate skill acquisition. Analysis of speed stacking time on a retention test revealed an overall benefit for the distributed relative to the massed practice group. Interestingly, our analysis of the benefits of distributed practice during training only showed performance benefits in the early session (session one) and later sessions (sessions eight, nine, and ten) of skill acquisition but not mid-way through it (sessions two through seven). Our results support previous findings highlighting the learning benefits of a distributed practice schedule but suggest that these benefits occur differentially throughout acquisition. Our work also replicates research demonstrating that observational learning is more beneficial when it is yoked to actual practice.
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http://dx.doi.org/10.1177/0031512520981242DOI Listing
April 2021

The ERP, frequency, and time-frequency correlates of feedback processing: Insights from a large sample study.

Psychophysiology 2021 02 10;58(2):e13722. Epub 2020 Nov 10.

Centre for Biomedical Research, University of Victoria, Victoria, BC, Canada.

Human learning, at least in part, appears to be dependent on the evaluation of how outcomes of our actions align with our expectations. Over the past 23 years, electroencephalography (EEG) has been used to probe the neural signatures of feedback processing. Seminal work demonstrated a difference in the human event-related potential (ERP) dependent on whether people were processing correct or incorrect feedback. Since then, these feedback evoked ERPs have been associated with reinforcement learning and conflict monitoring, tied to subsequent behavioral adaptations, and shown to be sensitive to a wide range of factors (e.g., Parkinson's disease). Recently, research has turned to frequency decomposition techniques to examine how changes in the EEG power spectra are related to underlying learning mechanisms. Although the literature on the neural correlates of feedback processing is vast, there are still methodological discrepancies and differences in results across studies. Here, we provide reference results and an investigation of methodological considerations for the ERP (reward positivity) and frequency (delta and theta power) correlates of feedback evaluation with a large sample size. Specifically, participants (n = 500) performed a two-armed bandit task while we recorded EEG. Our findings provide key information about the data characteristics and relationships that exist between the neural signatures of feedback evaluation. Additionally, we conclude with selected methodological recommendations for standardization of future research. All data and scripts are freely provided to facilitate open science.
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http://dx.doi.org/10.1111/psyp.13722DOI Listing
February 2021

Feedback processing is enhanced following exploration in continuous environments.

Neuropsychologia 2020 09 20;146:107538. Epub 2020 Jun 20.

Centre for Biomedical Research, University of Victoria, Victoria, British Columbia, V8W 2Y2, Canada.

Decision-making is typically studied by presenting participants with a small set of options. However, real-world behaviour, like foraging, often occurs in continuous environments. The degree to which human decision-making in discrete tasks generalizes to continuous tasks is questionable. For example, successful foraging comprises both exploration (learning about the environment) and exploitation (taking advantage of what is known). Although progress has been made in understanding the neural processes related to this trade-off in discrete tasks, it is currently unclear how, or whether, the same processes are involved in continuous tasks. To address this, we recorded electroencephalographic data while participants "dug for gold" by selecting locations on a map. Participants were cued beforehand that the map contained either a single patch of gold, or many patches of gold. We then used a computational model to classify participant responses as either exploitations, which were driven by previous reward locations and amounts, or explorations. Our participants were able to adjust their strategy based on reward distribution, exploring more in multi-patch environments and less in single-patch environments. We observed an enhancement of the feedback-locked P300, a neural signal previously linked to exploration in discrete tasks, which suggests the presence of a general neural system for managing the explore-exploit trade-off. Furthermore, the P300 was accompanied by an exploration-related enhancement of the late positive potential that was greatest in the multi-patch environment, suggesting a role for motivational processes during exploration.
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http://dx.doi.org/10.1016/j.neuropsychologia.2020.107538DOI Listing
September 2020

Visual Feedback Modulates Aftereffects and Electrophysiological Markers of Prism Adaptation.

Front Hum Neurosci 2020 17;14:138. Epub 2020 Apr 17.

Cognitive Health and Recovery Research Lab, Departments of Psychiatry, Psychology and Neuroscience, Brain Repair Centre, Dalhousie University, Halifax, NS, Canada.

Prism adaptation (PA) is both a model for visuomotor learning and a promising treatment for visuospatial neglect after stroke. The task involves reaching for targets while prism glasses horizontally displace the visual field. Adaptation is hypothesized to occur through two processes: strategic recalibration, a rapid self-correction of pointing errors; and spatial realignment, a more gradual adjustment of visuomotor reference frames that produce prism aftereffects (i.e., reaching errors upon glasses removal in the direction opposite to the visual shift). While aftereffects can ameliorate neglect, not all patients respond to PA, and the neural mechanisms underlying successful adaptation are unclear. We investigated the feedback-related negativity (FRN) and the P300 event-related potential (ERP) components as candidate markers of strategic recalibration and spatial realignment, respectively. Healthy young adults wore prism glasses and performed memory-guided reaching toward vertical-line targets. ERPs were recorded in response to three different between-subject error feedback conditions at screen-touch: view of hand and target (Experiment 1), view of hand only (Experiment 2), or view of lines to mark target and hand position (view of hand occluded; Experiment 3). Conditions involving a direct view of the hand-produced stronger aftereffects than indirect hand feedback, and also evoked a P300 that decreased in amplitude as adaptation proceeded. Conversely, the FRN was only seen in conditions involving target feedback, even when aftereffects were smaller. Since conditions producing stronger aftereffects were associated with a phase-sensitive P300, this component may index a "context-updating" realignment process critical for strong aftereffects, whereas the FRN may reflect an error monitoring process related to strategic recalibration.
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http://dx.doi.org/10.3389/fnhum.2020.00138DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7182100PMC
April 2020

Electroencephalographic evidence for a reinforcement learning advantage during motor skill acquisition.

Biol Psychol 2020 03 22;151:107849. Epub 2020 Jan 22.

Centre for Biomedical Research, University of Victoria, Canada. Electronic address:

The feedback that we receive shapes how we learn. Previous research has demonstrated that quantitative feedback results in better performance than qualitative feedback. However, the data supporting a quantitative feedback advantage are not conclusive and further little work has been done to examine the mechanistic neural differences that underlie the relative benefits of quantitative and qualitative feedback. To address these issues, participants learned a simple motor task in quantitative and qualitative feedback conditions while electroencephalographic (EEG) data were recorded. We found that participants were more accurate and had a larger neural response - the feedback related negativity - when qualitative feedback was provided. Our data suggest that qualitative feedback is more advantageous than quantitative feedback during the early stages of skill acquisition. Additionally, our findings support previous work suggesting that a reinforcement learning system within the human medial-frontal cortex plays a key role in motor skill acquisition.
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http://dx.doi.org/10.1016/j.biopsycho.2020.107849DOI Listing
March 2020

The impact of wellness on neural learning systems.

Neurosci Lett 2020 01 9;714:134537. Epub 2019 Oct 9.

Centre for Biomedical Research, University of Victoria, Victoria, BC, Canada.

Over the past 20 years there has been an increasing push for people to achieve or maintain "wellness" - a state in which one has not only physical but also mental and social well-being. While it may seem obvious that maintaining a state of wellness is beneficial, little research has been done to probe how maintaining a state of wellness impacts our brain. Here, we specifically examined the impact of wellness on a neural system within the medial-frontal cortex responsible for human reinforcement learning. Sixty-two undergraduate students completed the Perceived Wellness Survey after which they completed a computer-based learnable gambling game while electroencephalographic data were recorded. Within the game, participants were presented with a series of choices that either led to financial gains or losses. An analysis of our behavioral data indicated that participants were able to learn the underlying structure of the gambling game given that we observed improvements in performance. Concurrent with this, we observed an electroencephalographic response evoked by the evaluation of gambling outcomes - the reward positivity. Importantly, we found significant relationships between several aspects of wellness and the amplitude of the reward positivity. Given that the reward positivity is thought to reflect the function of a reinforcement learning system within the medial-frontal cortex, our results suggest that wellness impacts neural function - in this instance one of the systems responsible for human learning.
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http://dx.doi.org/10.1016/j.neulet.2019.134537DOI Listing
January 2020

Quantifying two-dimensional and three-dimensional stereoscopic learning in anatomy using electroencephalography.

NPJ Sci Learn 2019 17;4:10. Epub 2019 Jul 17.

1Department of Veterinary Clinical Diagnostic Sciences, Faculty of Veterinary Medicine, University of Calgary, Calgary, AB Canada.

Advances in computer visualization enabling both 2D and 3D representation have generated tools to aid perception of spatial relationships and provide a new forum for instructional design. A key knowledge gap is the lack of understanding of how the brain neurobiologically processes and learns from spatially presented content, and new quantitative variables are required to address this gap. The objective of this study was to apply quantitative neural measures derived from electroencephalography (EEG) to examine stereopsis in anatomy learning by comparing mean amplitude changes in N250 (related to object recognition) and reward positivity (related to responding to feedback) event related to potential components using a reinforcement-based learning paradigm. Health sciences students ( = 61) learned to identify and localize neuroanatomical structures using 2D, 3D, or a combination of models while EEG and behavioral (accuracy) data were recorded. Participants learning using 3D models had a greater object recognition (N250 amplitude) compared to those who learned from 2D models. Based on neurological results, interleaved learning incorporating both 2D and 3D models provided an advantage in learning, retention, and transfer activities represented by decreased reward positivity amplitude. Behavioral data did not have the same sensitivity as neural data for distinguishing differences in learning with and without stereopsis in these learning activities. Measuring neural activity reveals new insights in applied settings for educators to consider when incorporating stereoscopic models in the design of learning interventions.
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http://dx.doi.org/10.1038/s41539-019-0050-4DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6637108PMC
July 2019

The importance of agency in human reward processing.

Cogn Affect Behav Neurosci 2019 12;19(6):1458-1466

Centre for Biomedical Research, University of Victoria, P.O. Box 1700 STN CSC, Victoria, British Columbia, V8W 2Y2, Canada.

Converging evidence suggests that reinforcement learning (RL) signals exist within the human brain and that they play a role in the modification of behaviour. According to RL theory, prediction errors are used to update values associated with actions and/or predictive cues, thus facilitate decision-making. For example, the reward positivity-a feedback-sensitive component of the event-related brain potential (ERP)-is thought to index an RL prediction error. An unresolved question, however, is whether or not action is required to elicit a reward positivity. Reinforcement learning theory would predict that the reward positivity should diminish or disappear in the absence of action, but evidence for this claim is conflicting. To investigate the impact of cue, choice, and action on the amplitude of the reward positivity, we altered a two-armed bandit task by systematically removing these factors. The reward positivity was greatly reduced or absent in the altered versions of the task. This result highlights the key role of agency in producing learning signals, such as the reward positivity.
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http://dx.doi.org/10.3758/s13415-019-00730-2DOI Listing
December 2019

Ready, set, explore! Event-related potentials reveal the time-course of exploratory decisions.

Brain Res 2019 09 29;1719:183-193. Epub 2019 May 29.

Centre for Biomedical Research, University of Victoria, Victoria, British Columbia V8W 2Y2, Canada. Electronic address:

The decision trade-off between exploiting the known and exploring the unknown has been studied using a variety of approaches and techniques. Surprisingly, electroencephalography (EEG) has been underused in this area of study, even though its high temporal resolution has the potential to reveal the time-course of exploratory decisions. We addressed this issue by recording EEG data while participants tried to win as many points as possible in a two-choice gambling task called a two-armed bandit. After using a computational model to classify responses as either exploitations or explorations, we examined event-related potentials locked to two events preceding decisions to exploit/explore: the arrival of feedback, and the subsequent appearance of the next trial's choice stimuli. In particular, we examined the feedback-locked P300 component, thought to index a phasic release of norepinephrine (a neural interrupt signal), and the reward positivity, thought to index a phasic release of dopamine (a neural prediction error signal). We observed an exploration-dependent enhancement of the P300 only, suggesting a critical role of norepinephrine (but not dopamine) in triggering decisions to explore. Similarly, we examined the N200/P300 components evoked by the appearance of the choice stimuli. In this case, exploration was characterized by an enhancement of the N200, but not P300, a result we attribute to increased response conflict. These results demonstrate the usefulness of combining computational and EEG methodologies, and suggest that exploratory decisions are preceded by two characterizing events: a feedback-locked neural interrupt signal (enhanced P300), and a choice-locked increase in response conflict (enhanced N200).
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http://dx.doi.org/10.1016/j.brainres.2019.05.039DOI Listing
September 2019

Passively learned spatial navigation cues evoke reinforcement learning reward signals.

Cognition 2019 08 28;189:65-75. Epub 2019 Mar 28.

Centre for Biomedical Research, University of Victoria, Canada.

Since the suggestion by Tolman (1948) that both rodents and humans create cognitive maps during navigation, the specifics of how navigators learn about their environment has been mired in debate. One facet of this debate is whether or not the creation of cognitive maps - also known as allocentric navigation - involves reinforcement learning. Here, we demonstrate a role for reinforcement learning during allocentric navigation using event-related brain potentials (ERPs). In the present experiment, participants navigated in a virtual environment that allowed the use of three different navigation strategies (allocentric, egocentric-response, & egocentric-cue), in which their goal was to locate and remember a hidden platform. Following the navigation phase of the experiment, participants were shown "cue images" representative of the three navigation strategies. Specifically, we examined whether or not these passively learned strategy images elicited a reward positivity - an ERP component associated with reinforcement learning and the anterior cingulate cortex. We found that when allocentric navigators were shown previously learned cues predicting the goal location a reward positivity was elicited. The present findings demonstrate that allocentric navigational cues carry long-term value after navigation and lend support to the claim that reinforcement learning plays a role in the acquisition of allocentric navigation and thus the generation of cognitive maps.
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http://dx.doi.org/10.1016/j.cognition.2019.03.015DOI Listing
August 2019

Thinking theta and alpha: Mechanisms of intuitive and analytical reasoning.

Neuroimage 2019 04 22;189:574-580. Epub 2019 Jan 22.

Centre for Biomedical Research, University of Victoria, 3800 Finnerty Rd, Victoria, British Columbia, V8P 5C2, Canada.

Humans have a unique ability to engage in different modes of thinking. Intuitive thinking (coined System 1, see Kahneman, 2011) is fast, automatic, and effortless whereas analytical thinking (coined System 2) is slow, contemplative, and effortful. We extend seminal pupillometry research examining these modes of thinking by using electroencephalography (EEG) to decipher their respective underlying neural mechanisms. We demonstrate that System 1 thinking is characterized by an increase in parietal alpha EEG power reflecting autonomic access to long-term memory and a release of attentional resources whereas System 2 thinking is characterized by an increase in frontal theta EEG power indicative of the engagement of cognitive control and working memory processes. Consider our results in terms of an example - a child may need cognitive control and working memory when contemplating a mathematics problem yet an adult can drive a car with little to no attention by drawing on easily accessed memories. Importantly, the unravelling of intuitive and analytical thinking mechanisms and their neural signatures will provide insight as to how different modes of thinking drive our everyday lives.
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http://dx.doi.org/10.1016/j.neuroimage.2019.01.048DOI Listing
April 2019

High-intensity interval exercise impairs neuroelectric indices of reinforcement-learning.

Physiol Behav 2019 01 5;198:18-26. Epub 2018 Oct 5.

School of Kinesiology and Health Studies, Queen's University, Kingston, ON K7L 3N6, Canada. Electronic address:

A single bout of high-intensity interval exercise (HIIE) improves behavioural measures of cognitive function; however, investigations using event-related potentials (ERPs) to examine the systems that underlie these cognitive improvements are lacking. The reward positivity is a positive-going ERP component that indexes reward processing evoked by 'win' feedback and is a candidate marker of an underlying human reinforcement learning system. While HIIE improves behavioural measures of learning, it is unknown how HIIE affects the amplitude of the reward positivity. Therefore, the purpose of this study was to investigate how HIIE affects reward positivity amplitude in response to reward feedback in university students. Using a single-group randomly assigned counterbalance crossover design, 25 healthy university students performed HIIE and control visits on separate days. Electroencephalographic data was recorded before (pre-intervention) and 10 min after (post-intervention) the intervention period while participants played a novel gambling task. The HIIE intervention consisted of 4 separate body-weight exercises totaling 11 min in duration, including rest. The control visit intervention consisted of quietly watching a nature documentary for 11 min. Heart rate (HR) was measured at the same time intervals in both trials. Analysis revealed that HIIE significantly diminished the amplitude of the reward positivity whereas it remained unaffected in the control condition. HR was significantly higher following HIIE compared to control during post-intervention testing. These findings suggest that mechanisms of reinforcement learning are impaired shortly after HIIE cessation, possibly due to persistent, suboptimal arousal as evidenced by elevated HR post-HIIE.
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http://dx.doi.org/10.1016/j.physbeh.2018.10.005DOI Listing
January 2019

Chasing the zone: Reduced beta power predicts baseball batting performance.

Neurosci Lett 2018 11 6;686:150-154. Epub 2018 Sep 6.

Centre for Biomedical Research, University of Victoria, Victoria, B.C., Canada. Electronic address:

Mental state prior to sports skill execution is related to subsequent performance. For example, relationships between pre-performance electroencephalogram (EEG) power and subsequent movement outcomes in golf putting, pistol shooting, and basketball free throw shooting have been previously reported. With that said, the existing body of research examining the pre-performance EEG - performance relationship has been focused on the execution of internally as opposed to externally-paced motor skills. Given that the execution of internally and externally-paced movements are dependent on different neural pathways, in the present study we examined whether or not pre-performance EEG power predicted ensuing performance of an externally-paced motor skill - baseball batting. Sixty-seven baseball players had EEG data recorded for 120 s prior to batting practice. Performance was assessed by three expert coaches and the accuracy of coach performance ratings was verified via Generalizability Theory. An analysis of our data revealed an inverse relationship between frontal EEG power in the beta range and subsequent batting performance - reduced beta power was associated with better batting performance whereas increased beta power was associated with worse batting performance. Our results are in line with prior research that has demonstrated a relationship between increased EEG power in the beta range and the subsequent commitment of motor errors in addition to the aforementioned work examining pre-performance EEG and the execution of internally-paced motor skills.
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http://dx.doi.org/10.1016/j.neulet.2018.09.004DOI Listing
November 2018

Alcohol hangover impacts learning and reward processing within the medial-frontal cortex.

Psychophysiology 2018 08 30;55(8):e13081. Epub 2018 Mar 30.

Centre for Biomedical Research, University of Victoria, Victoria, British Columbia, Canada.

It is common knowledge that alcohol intoxication impairs motor coordination, judgment, and decision making. Indeed, an abundance of literature links intoxication to impaired cognitive control that leads to accidents and injury. A broadening body of research, however, suggests that the impact of alcohol may continue beyond the point of intoxication and into the period of alcohol hangover. Here, we examined differences in the amplitude of reward positivity-a component of the human ERP associated with learning-between control and hangover participants. During performance of a learnable gambling task, we found a reduction in the reward positivity during alcohol hangover. Additionally, participants experiencing alcohol hangover demonstrated reduced performance in the experimental task in comparison to their nonhangover counterparts. Our results suggest that the neural systems that underlie performance monitoring and reward-based learning are impaired during alcohol hangover.
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http://dx.doi.org/10.1111/psyp.13081DOI Listing
August 2018

Learning what matters: A neural explanation for the sparsity bias.

Int J Psychophysiol 2018 05 15;127:62-72. Epub 2018 Mar 15.

Centre for Biomedical Research, University of Victoria, Victoria, British Columbia V8W 2Y2, Canada.

The visual environment is filled with complex, multi-dimensional objects that vary in their value to an observer's current goals. When faced with multi-dimensional stimuli, humans may rely on biases to learn to select those objects that are most valuable to the task at hand. Here, we show that decision making in a complex task is guided by the sparsity bias: the focusing of attention on a subset of available features. Participants completed a gambling task in which they selected complex stimuli that varied randomly along three dimensions: shape, color, and texture. Each dimension comprised three features (e.g., color: red, green, yellow). Only one dimension was relevant in each block (e.g., color), and a randomly-chosen value ranking determined outcome probabilities (e.g., green > yellow > red). Participants were faster to respond to infrequent probe stimuli that appeared unexpectedly within stimuli that possessed a more valuable feature than to probes appearing within stimuli possessing a less valuable feature. Event-related brain potentials recorded during the task provided a neurophysiological explanation for sparsity as a learning-dependent increase in optimal attentional performance (as measured by the N2pc component of the human event-related potential) and a concomitant learning-dependent decrease in prediction errors (as measured by the feedback-elicited reward positivity). Together, our results suggest that the sparsity bias guides human reinforcement learning in complex environments.
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http://dx.doi.org/10.1016/j.ijpsycho.2018.03.006DOI Listing
May 2018

A Reinforcement-Based Learning Paradigm Increases Anatomical Learning and Retention-A Neuroeducation Study.

Front Hum Neurosci 2018 6;12:38. Epub 2018 Feb 6.

Department of Cell Biology and Anatomy, Cumming School of Medicine, University of Calgary, Calgary, AB, Canada.

In anatomy education, a key hurdle to engaging in higher-level discussion in the classroom is recognizing and understanding the extensive terminology used to identify and describe anatomical structures. Given the time-limited classroom environment, seeking methods to impart this foundational knowledge to students in an efficient manner is essential. Just-in-Time Teaching (JiTT) methods incorporate pre-class exercises (typically online) meant to establish foundational knowledge in novice learners so subsequent instructor-led sessions can focus on deeper, more complex concepts. Determining how best do we design and assess pre-class exercises requires a detailed examination of learning and retention in an applied educational context. Here we used electroencephalography (EEG) as a quantitative dependent variable to track learning and examine the efficacy of JiTT activities to teach anatomy. Specifically, we examined changes in the amplitude of the N250 and reward positivity event-related brain potential (ERP) components alongside behavioral performance as novice students participated in a series of computerized reinforcement-based learning modules to teach neuroanatomical structures. We found that as students learned to identify anatomical structures, the amplitude of the N250 increased and reward positivity amplitude decreased in response to positive feedback. Both on a retention and transfer exercise when learners successfully remembered and translated their knowledge to novel images, the amplitude of the reward positivity remained decreased compared to early learning. Our findings suggest ERPs can be used as a tool to track learning, retention, and transfer of knowledge and that employing the reinforcement learning paradigm is an effective educational approach for developing anatomical expertise.
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http://dx.doi.org/10.3389/fnhum.2018.00038DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5808130PMC
February 2018

Visuomotor mental rotation of a saccade: The contingent negative variation scales to the angle of rotation.

Vision Res 2018 02 19;143:82-88. Epub 2017 Dec 19.

Neuroeducation Network, University of Victoria, Victoria, BC, Canada.

The visuomotor mental rotation (VMR) of a saccade requires a response to a region of space that is dissociated from a stimulus by a pre-specified angle, and work has shown a monotonic increase in reaction times as a function of increasing oblique angles of rotation. These results have been taken as evidence of a continuous process of rotation and have generated competing hypotheses. One hypothesis asserts that rotation is mediated via frontoparietal structures, whereas a second states that a continuous shift in the activity of direction-specific neurons in the superior colliculus (SC) supports rotation. Research to date, however, has not examined the neural mechanisms underlying VMR saccades and both hypotheses therefore remain untested. The present study measured the behavioural data and event-related brain potentials (ERP) of standard (i.e., 0° of rotation) and VMR saccades involving 35°, 70° and 105° of rotation. Behavioural results showed that participants adhered to task-based rotation demands and ERP findings showed that the amplitude of the contingent negative variation (CNV) linearly decreased with increasing angle of rotation. The cortical generators of the CNV are linked to frontoparietal structures supporting movement preparation. Although our ERP design does not allow us to exclude a possible role of the SC in the rotation of a VMR saccade, they do demonstrate that such actions are supported by a continuous and cortically based rotation process.
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http://dx.doi.org/10.1016/j.visres.2017.09.005DOI Listing
February 2018

Event-related brain potentials and the study of reward processing: Methodological considerations.

Int J Psychophysiol 2018 10 14;132(Pt B):175-183. Epub 2017 Nov 14.

Centre for Biomedical Research, University of Victoria, P.O. Box 17000 STN CSC, Victoria, British Columbia V8W 2Y2, Canada. Electronic address:

There is growing interest in using electroencephalography and specifically the event-related brain potential (ERP) methodology to study human reward processing. Since the discovery of the feedback related negativity (Miltner et al., 1997) and the development of theories associating the feedback related negativity and more recently the reward positivity with reinforcement learning, midbrain dopamine function, and the anterior cingulate cortex (i.e., Holroyd and Coles, 2002) researchers have used the ERP methodology to probe the neural basis of reward learning in humans. However, examination of the feedback related negativity and the reward positivity cannot be done without an understanding of some key methodological issues that must be taken into account when using ERPs and examining these ERP components. For example, even the component name - the feedback related negativity - is a source of debate within the research community as some now strongly feel that the component should be named the reward positivity (Proudfit, 2015). Here, ten key methodological issues are discussed - confusion in component naming, the reward positivity, component identification, peak quantification and the use of difference waveforms, frequency (the N200) and component contamination (the P300), the impact of feedback timing, action, and task learnability, and how learning results in changes in the amplitude of the feedback-related negativity/reward positivity. The hope here is to not provide a definitive approach for examining the feedback related negativity/reward positivity, but instead to outline the key issues that must be taken into account when examining this component to assist researchers in their study of human reward processing with the ERP methodology.
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http://dx.doi.org/10.1016/j.ijpsycho.2017.11.007DOI Listing
October 2018

The application of reward learning in the real world: Changes in the reward positivity amplitude reflect learning in a medical education context.

Int J Psychophysiol 2018 10 27;132(Pt B):236-242. Epub 2017 Oct 27.

Centre for Biomedical Research, University of Victoria, Canada.

Evidence ranging from behavioural adaptations to neurocognitive theories has made significant advances into our understanding of feedback-based learning. For instance, over the past twenty years research using electroencephalography has demonstrated that the amplitude of a component of the human event-related brain potential - the reward positivity - appears to change with learning in a manner predicted by reinforcement learning theory (Holroyd and Coles, 2002; Sutton and Barto, 1998). However, while the reward positivity (also known as the feedback related negativity) is well studied, whether the component reflects an underlying learning process or whether it is simply sensitive to feedback evaluation is still unclear. Here, we sought to provide support that the reward positivity is reflective of an underlying learning process and further we hoped to demonstrate this in a real-world medical education context. In the present study, students with no medical training viewed a series of patient cards that contained ten physiological readings relevant for diagnosing liver and biliary disease types, selected the most appropriate diagnostic classification, and received feedback as to whether their decisions were correct or incorrect. Our behavioural results revealed that our participants were able to learn to diagnose liver and biliary disease types. Importantly, we found that the amplitude of the reward positivity diminished in a concomitant manner with the aforementioned behavioural improvements. In sum, our data support theoretical predictions (e.g., Holroyd and Coles, 2002), suggest that the reward positivity is an index of a neural learning system, and further validate that this same system is involved in learning across a wide range of contexts.
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http://dx.doi.org/10.1016/j.ijpsycho.2017.10.010DOI Listing
October 2018

When theory and biology differ: The relationship between reward prediction errors and expectancy.

Biol Psychol 2017 10 18;129:265-272. Epub 2017 Sep 18.

Centre for Biomedical Research, University of Victoria, Victoria, British Columbia, V8W 2Y2, Canada.

Comparisons between expectations and outcomes are critical for learning. Termed prediction errors, the violations of expectancy that occur when outcomes differ from expectations are used to modify value and shape behaviour. In the present study, we examined how a wide range of expectancy violations impacted neural signals associated with feedback processing. Participants performed a time estimation task in which they had to guess the duration of one second while their electroencephalogram was recorded. In a key manipulation, we varied task difficulty across the experiment to create a range of different feedback expectancies - reward feedback was either very expected, expected, 50/50, unexpected, or very unexpected. As predicted, the amplitude of the reward positivity, a component of the human event-related brain potential associated with feedback processing, scaled inversely with expectancy (e.g., unexpected feedback yielded a larger reward positivity than expected feedback). Interestingly, the scaling of the reward positivity to outcome expectancy was not linear as would be predicted by some theoretical models. Specifically, we found that the amplitude of the reward positivity was about equivalent for very expected and expected feedback, and for very unexpected and unexpected feedback. As such, our results demonstrate a sigmoidal relationship between reward expectancy and the amplitude of the reward positivity, with interesting implications for theories of reinforcement learning.
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http://dx.doi.org/10.1016/j.biopsycho.2017.09.007DOI Listing
October 2017

Older adults display diminished error processing and response in a continuous tracking task.

Psychophysiology 2017 Nov 16;54(11):1706-1713. Epub 2017 Jun 16.

Centre for Biomedical Research, University of Victoria, Victoria, British Columbia, Canada.

Advancing age is often accompanied by a decline in motor control that results in a decreased ability to successfully perform motor tasks. While there are multiple factors that contribute to age-related deficits in motor control, one unexplored possibility is that age-related deficits in our ability to evaluate motor output result in an increase in motor errors. In line with this, previous work from our laboratory demonstrated that motor errors evoked an error-related negativity (ERN)-a component of the human ERP associated with error evaluation originating within the human medial-frontal cortex. In the present study, we examined whether or not deficits in the medial-frontal error evaluation system contribute to age-related deficits in motor control. Two groups of participants (young, old) performed a computer-based tracking task that paralleled driving while EEG data were recorded. Our results show that older adults committed more behavioral errors than young adults during performance of the tracking task. An analysis of our ERP data revealed that the amplitude of the ERN was reduced in older adults relative to young adults following motor errors. Our results make an important extension from previous work demonstrating age-related reductions in the ERN during performance of cognitive tasks. Importantly, our results imply the possibility of understanding motor deficits in older age.
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http://dx.doi.org/10.1111/psyp.12907DOI Listing
November 2017

Choosing MUSE: Validation of a Low-Cost, Portable EEG System for ERP Research.

Front Neurosci 2017 10;11:109. Epub 2017 Mar 10.

Neuroeconomics Laboratory, Centre for Biomedical Research, University of Victoria Victoria, BC, Canada.

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 -tests of component existence (all '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.
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http://dx.doi.org/10.3389/fnins.2017.00109DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5344886PMC
March 2017

A preliminary investigation into the neural basis of the production effect.

Can J Exp Psychol 2016 Jun;70(2):139-46

Neuroeconomics Laboratory, University of Victoria.

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
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http://dx.doi.org/10.1037/cep0000093DOI Listing
June 2016

The scarcity heuristic impacts reward processing within the medial-frontal cortex.

Neuroreport 2016 May;27(7):522-6

aNeuroeconomics Laboratory, Department of Exercise Science, Physical & Health Education, University of Victoria, Victoria bDepartment of Psychology, University of British Columbia, Vancouver, British Columbia, Canada.

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.
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http://dx.doi.org/10.1097/WNR.0000000000000575DOI Listing
May 2016
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