Contribution of the sensorimotor beta oscillations and the cortico-basal ganglia-thalamic circuitry during value-based decision making: A simultaneous EEG-fMRI investigation.

In decision neuroscience, the motor system has primarily been considered to be involved in executing choice actions. However, a competing perspective suggests its engagement in the evaluation of options, traditionally considered to be performed by the brain's valuation system. Here, we investigate the role of the motor system in value-based decision making by determining the neural circuitries associated with the sensorimotor beta oscillations previously identified to encode decision options. In a simultaneous EEG-fMRI study, participants evaluated reward and risk associated with a forthcoming action. A significant sensorimotor beta desynchronization was identified prior to and independent of response. The level of beta desynchronization showed evidence of encoding the reward levels. This beta desynchronization covaried, on a trial-by-trial level, with BOLD activity in the cortico-basal ganglia-thalamic circuitry. In contrast, there was only a weak covariation within the valuation network, despite significant modulation of its BOLD activity by reward levels. These results suggest that the way in which decision variables are processed differs in the valuation network and in the cortico-basal ganglia-thalamic circuitry. We propose that sensorimotor beta oscillations indicate incentive motivational drive towards a choice action computed from the decision variables even prior to making a response, and it arises from the cortico-basal ganglia-thalamic circuitry.

Reward and expectancy effects on neural signals of motor preparation and execution.

The magnitude and prospect of rewards can have strong modulatory effects on response preparation and execution. Importantly, reward-seeking behavior in real life happens under an environment characterized by uncertainty and dynamic changes. The current study investigated how the brain's motor and cognitive control system adapts to the dynamic changes in the environment in pursuit of rewards. To this end, we tested the effect of reward and expectancy on the hallmark neural signals that reflect activity in motor and prefrontal systems, the lateralized readiness potential (LRP) and the mediofrontal (mPFC) theta oscillations, while participants performed an expected and unexpected actions to retrieve rewards. To better capture the dynamic changes in neural processes represented in the LRP waveform, we decomposed the LRP into the preparation (LRPprep) and execution (LRPexec) components. In the comparison of LRP magnitude across task conditions, we found a greater LRPprep when large rewards were more likely, reflecting a greater motor preparation to obtain larger rewards. We also found a greater LRPexec when large rewards were presented unexpectedly, suggesting a greater motor effort placed for executing a correct movement when presented with large rewards. In the analysis of mPFC theta, we found a greater theta power prior to performing an unexpected than expected response, indicating its contribution in response conflict resolution. Collectively, these results demonstrate an optimized motor and cognitive control to maximize rewards under the dynamic changes of real-life environment.

Cultural modulation of early attentional responses to positive self-information: An ERP investigation of self-enhancement.

Westerners show a strong tendency to view themselves in a positive light (i.e., self-enhancement), but this tendency is substantially weaker among East Asians. At present, however, it remains inconclusive whether this cultural difference reflects genuine variation in spontaneous engagement in this motivational tendency or is driven by individuals' deliberate efforts to present oneself in culturally acceptable ways. In the present research, we sought to address this issue by examining whether culture modulates early attentional bias to self-name, presented in the context of positive (vs. negative) situations. Both European American and Asian American participants (N = 64) read a series of scenarios depicting either a positive or a negative life situation and were subsequently presented with their name or names of famous people or strangers. European Americans showed greater P2, an ERP component of early attentional arousal, when the self-name was primed with positive (vs. negative) situations. In contrast, Asian Americans' P2 in response to the self-name was not moderated by situation primes. Moreover, the degree to which P2 was enhanced in positive (vs. negative) situations was negatively correlated with Asian cultural values. These results demonstrate that cultural variation in self-enhancement may reflect genuine beliefs about the self, manifested automatically at an early stage of attentional processing.

Neurocognitive underpinnings of cross-cultural differences in risky decision making.

Culture permeates across human mind and behavior. Cultural influence is reported even in economic decision making, which involves basic cognitive process, once believed to be invariant across all humans. The current study investigated the neurocognitive processes underlying economic decision making in East Asians and European Americans, with an aim to understand the cross-cultural differences in the discrete mental processes of decision making. Participants performed a risky gambling task that captures the gain maximizing and loss minimizing strategies, while electroencephalography was simultaneously collected. Event-related potentials (ERPs) associated with spontaneous emotional arousal (P2) and effortful attentional allocation (P3) were examined to determine the cultural effects on mental processes during pre-decisional and post-decisional stages. Behaviorally, Americans showed greater loss minimization than Asians. ERPs demonstrated significant cultural differences during post-decisional evaluation of outcomes, but not during pre-decisional processes. In Asians', ERP associated with emotional arousal (P2) was strongly modulated by gains, while in Americans', ERP associated with attentional allocation (P3) was strongly modulated by losses. These results suggest that Americans make conscious efforts to be self-reliant when facing financial losses, whereas Asians are more emotionally aroused by financial gains, which invites a refinement to the current theoretical propositions about cultural influence on decision making.

Money for me and money for friend: An ERP study of social reward processing in adolescents and adults.

Peer relations during adolescence contribute significantly to the development of socio-cognitive skills and pro-sociality. The current study probed the characteristics of adolescent socio-cognitive processing through a card game where they earn money for self and friend. We investigated the choice preference and temporal dynamics of information processing by measuring ERP responses to wins or losses (valence) directed toward self and friend (recipient). Choice data showed that despite adults and adolescents earning equivalent amounts across recipients combined, adults won significantly more for self than a friend; no such difference was found in adolescents. The ERPs in response to choice outcomes showed that the valence information was processed earlier (at P2) in adults, while it was processed later (at P3) in adolescents. Furthermore, a strong effect of recipient was present in adults later in the time course (at P3), while such an effect was weak in adolescents; if any, adolescents showed sensitivity to recipient information earlier at P2. These ERP data suggest a relatively equal allocation of the P3-mediated attentional process to both self and friend's outcomes in adolescents, which parallels the choice behavior. Collectively our results characterize adolescent pro-sociality toward friends, reflecting the importance of peer relationship during this unique developmental period.

"Incentive hope" and the nature of impulsivity in low-socioeconomic-status individuals.

Low-income environments have been associated with greater levels of impulsive behavior, which contribute to the higher debt and obesity rates that further perpetuate current wealth and health disparities. In this commentary, we describe how this might be explained by an appeal to "incentive hope" and the motivational drive toward consumption triggered by the future uncertainty these groups face.

Contribution of sensorimotor beta oscillations during value-based action selection.

Choosing a course of daily-life actions requires an accurate assessment of the associated risks and potential rewards. We investigated the neural dynamics of this decision process by analyzing the neural electrical signals acquired from electroencephalography (EEG) during a value-based action-selection task. In particular, we determined whether sensorimotor beta oscillations, traditionally studied in the context of motor control, are also involved in value-based decision making for actions. Additionally, we examined the involvement of this beta signal relative to other neural signals such as the ERP components P2 and P3, which have been previously identified in reward processing and value computations. Our results from healthy young adults (N = 31), showed a significant decrease in sensorimotor beta power during a decision phase without any motor response, in addition to an action phase when a response was made. The decision-phase beta signal was preceded by the P2/P3b components, and all of these neural signals reliably dissociated the different reward and risk levels, suggesting the encoding of decision variables. Importantly, while the beta signal during both the action and decision phase predicted behavioral performance (i.e., response time) in the action phase, the preceding P2/P3b had no such predictive association with the behavior. Collectively, these results demonstrate a unique contribution of the motor system in value-based decision making for actions, via the translation of motivational information into a motor signal across time.

The order of information processing alters economic gain-loss framing effects.

Adaptive decision making requires analysis of available information during the process of choice. In many decisions that information is presented visually - which means that variations in visual properties (e.g., salience, complexity) can potentially influence the process of choice. In the current study, we demonstrate that variation in the left-right positioning of risky and safe decision options can influence the canonical gain-loss framing effect. Two experiments were conducted using an economic framing task in which participants chose between gambles and certain outcomes. The first experiment demonstrated that the magnitude of the gain-loss framing effect was greater when the certain option signaling the current frame was presented on the left side of the visual display. Eye-tracking data during task performance showed a left-gaze bias for initial fixations, suggesting that the option presented on the left side was processed first. Combination of eye-tracking and choice data revealed that there was a significant effect of direction of first gaze (i.e. left vs. right) as well as an interaction between gaze direction and identity of the first fixated information (i.e. certain vs. gamble) regardless of frame. A second experiment presented the gamble and certain options in a random order, with a temporal delay between their presentations. We found that the magnitude of gain-loss framing was larger when the certain option was presented first, regardless of left and right positioning, only in individuals with lower risk-taking tendencies. The effect of presentation order on framing was not present in high risk-takers. These results suggest that the sequence of visual information processing as well as their left-right positioning can bias choices by changing the impact of the presented information during risky decision making.

What Makes You Go Faster?: The Effect of Reward on Speeded Action under Risk.

Evaluating the potential reward and risk associated with a choice of action plays an important role in everyday decision making. However, the details behind how reward and risk affect the decisions for actions remain unclear. The present study investigates the influence of reward and risk on a decision to make a speeded motor response. One hundred and ten college students performed a Speed-Rewarded Go-NoGo task during which they were rewarded proportionally based on the speed and accuracy of their response. On each trial, the magnitude of potential reward and the probability of a forthcoming Go signal (Go-probability) were presented prior to the Go or NoGo signal. Personality traits, such as risk taking and impulsive tendencies, were measured to determine their contribution in explaining individual differences in task performance. The results showed that larger amount of rewards can motivate people to respond faster, and this effect was modulated by the assessed risk, suggesting that decisions for actions are based on a systematic trade-off between rewards and risks. Moreover, when the assessed risk was high, individuals with greater risk taking and impulsive tendencies did not adequately adjust their behavior across different reward levels. These findings shed light on the mechanistic understanding of the effect of reward and risk on decisions for a speeded action.

Prosocial Reward Learning in Children and Adolescents.

Adolescence is a period of increased sensitivity to social contexts. To evaluate how social context sensitivity changes over development-and influences reward learning-we investigated how children and adolescents perceive and integrate rewards for oneself and others during a dynamic risky decision-making task. Children and adolescents (N = 75, 8-16 years) performed the Social Gambling Task (SGT, Kwak et al., 2014) and completed a set of questionnaires measuring other-regarding behavior. In the SGT, participants choose amongst four card decks that have different payout structures for oneself and for a charity. We examined patterns of choices, overall decision strategies, and how reward outcomes led to trial-by-trial adjustments in behavior, as estimated using a reinforcement-learning model. Performance of children and adolescents was compared to data from a previously collected sample of adults (N = 102) performing the identical task. We found that that children/adolescents were not only more sensitive to rewards directed to the charity than self but also showed greater prosocial tendencies on independent measures of other-regarding behavior. Children and adolescents also showed less use of a strategy that prioritizes rewards for self at the expense of rewards for others. These results support the conclusion that, compared to adults, children and adolescents show greater sensitivity to outcomes for others when making decisions and learning about potential rewards.

Altruistic traits are predicted by neural responses to monetary outcomes for self vs charity.

Human altruism is often expressed through charitable donation-supporting a cause that benefits others in society, at cost to oneself. The underlying mechanisms of this other-regarding behavior remain imperfectly understood. By recording event-related-potential (ERP) measures of brain activity from human participants during a social gambling task, we identified markers of differential responses to receipt of monetary outcomes for oneself vs for a charitable cause. We focused our ERP analyses on the frontocentral feedback-related negativity (FRN) and three subcomponents of the attention-related P300 (P3) brain wave: the frontocentral P2 and P3a and the parietal P3b. The FRN distinguished between gains and losses for both self and charity outcomes. Importantly, this effect of outcome valence was greater for self than charity for both groups and was independent of two altruism-related measures: participants' pre-declared intended donations and the actual donations resulting from their choices. In contrast, differences in P3 subcomponents for outcomes for self vs charity strongly predicted both of our laboratory measures of altruism-as well as self-reported engagement in real-life altruistic behaviors. These results indicate that individual differences in altruism are linked to individual differences in the relative deployment of attention (as indexed by the P3) toward outcomes affecting other people.

Interactive effects of age and multi-gene profile on motor learning and sensorimotor adaptation.

The interactive association of age and dopaminergic polymorphisms on cognitive function has been studied extensively. However, there is limited research on whether age interacts with the association between genetic polymorphisms and motor learning. We examined a group of young and older adults' performance in three motor tasks: explicit sequence learning, visuomotor adaptation, and grooved pegboard. We assessed whether individuals' motor learning and performance were associated with their age and genotypes. We selected three genetic polymorphisms: Catechol-O-Methyl Transferase (COMT val158met) and Dopamine D2 Receptor (DRD2 G>T), which are involved with dopaminergic regulation, and Brain Derived Neurotrophic Factor (BDNF val66met) that modulates neuroplasticity and has been shown to interact with dopaminergic genes. Although the underlying mechanisms of the function of these three genotypes are different, the high performance alleles of each have been linked to better learning and performance. We created a composite polygene score based on the Number of High Performance Alleles (NHPA) that each individual carried. We found several associations between genetic profile, motor performance, and sensorimotor adaptation. More importantly, we found that this association varies with age, task type, and engagement of implicit versus explicit learning processes.

The Rational Adolescent: Strategic Information Processing during Decision Making Revealed by Eye Tracking.

Adolescence is often viewed as a time of irrational, risky decision-making - despite adolescents' competence in other cognitive domains. In this study, we examined the strategies used by adolescents (N=30) and young adults (N=47) to resolve complex, multi-outcome economic gambles. Compared to adults, adolescents were more likely to make conservative, loss-minimizing choices consistent with economic models. Eye-tracking data showed that prior to decisions, adolescents acquired more information in a more thorough manner; that is, they engaged in a more analytic processing strategy indicative of trade-offs between decision variables. In contrast, young adults' decisions were more consistent with heuristics that simplified the decision problem, at the expense of analytic precision. Collectively, these results demonstrate a counter-intuitive developmental transition in economic decision making: adolescents' decisions are more consistent with rational-choice models, while young adults more readily engage task-appropriate heuristics.

Measurement of Shoulder Range of Motion in Patients with Adhesive Capsulitis Using a Kinect.

Range of motion (ROM) measurements are essential for the evaluation for and diagnosis of adhesive capsulitis of the shoulder (AC). However, taking these measurements using a goniometer is inconvenient and sometimes unreliable. The Kinect (Microsoft, Seattle, WA, USA) is gaining attention as a new motion detecting device that is nonintrusive and easy to implement. This study aimed to apply Kinect to measure shoulder ROM in AC; we evaluated its validity by calculating the agreement of the measurements obtained using Kinect with those obtained using goniometer and assessed its utility for the diagnosis of AC. Both shoulders of 15 healthy volunteers and affected shoulders of 12 patients with AC were included in the study. The passive and active ROM of each were measured with a goniometer for flexion, abduction, and external rotation. Their active shoulder motions for each direction were again captured using Kinect and the ROM values were calculated. The agreement between the two measurements was tested with the intraclass correlation coefficient (ICC). Diagnostic performance using the Kinect ROM was evaluated with Cohen's kappa value. The cutoff values of the limited ROM were determined in the following ways: the same as passive ROM values, reflecting the mean difference, and based on receiver operating characteristic curves. The ICC for flexion/abduction/external rotation between goniometric passive ROM and the Kinect ROM were 0.906/0.942/0.911, while those between active ROMs and the Kinect ROMs were 0.864/0.932/0.925. Cohen's kappa values were 0.88, 0.88, and 1.0 with the cutoff values in the order above. Measurements of the shoulder ROM using Kinect show excellent agreement with those taken using a goniometer. These results indicate that the Kinect can be used to measure shoulder ROM and to diagnose AC as an alternative to goniometer.

Altered cerebellar connectivity in Parkinson's patients ON and OFF L-DOPA medication.

Although nigrostriatal changes are most commonly affiliated with Parkinson's disease, the role of the cerebellum in Parkinson's has become increasingly apparent. The present study used lobule-based cerebellar resting state functional connectivity to (1) compare cerebellar-whole brain and cerebellar-cerebellar connectivity in Parkinson's patients both ON and OFF L-DOPA medication and controls, and to (2) relate variations in cerebellar connectivity to behavioral performance. Results indicated that, when contrasted to the control group, Parkinson's patients OFF medication had increased levels of cerebellar-whole brain and cerebellar-cerebellar connectivity, whereas Parkinson's patients ON medication had decreased levels of cerebellar-whole brain and cerebellar-cerebellar connectivity. Moreover, analyses relating levels of cerebellar connectivity to behavioral measures demonstrated that, within each group, increased levels of connectivity were most often associated with improved cognitive and motor performance, but there were several instances where increased connectivity was related to poorer performance. Overall, the present study found medication-variant cerebellar connectivity in Parkinson's patients, further demonstrating cerebellar changes associated with Parkinson's disease and the moderating effects of medication.

Differential reward learning for self and others predicts self-reported altruism.

In social environments, decisions not only determine rewards for oneself but also for others. However, individual differences in pro-social behaviors have been typically studied through self-report. We developed a decision-making paradigm in which participants chose from card decks with differing rewards for themselves and charity; some decks gave similar rewards to both, while others gave higher rewards for one or the other. We used a reinforcement-learning model that estimated each participant's relative weighting of self versus charity reward. As shown both in choices and model parameters, individuals who showed relatively better learning of rewards for charity--compared to themselves--were more likely to engage in pro-social behavior outside of a laboratory setting indicated by self-report. Overall rates of reward learning, however, did not predict individual differences in pro-social tendencies. These results support the idea that biases toward learning about social rewards are associated with one's altruistic tendencies.

Lifespan differences in cortico-striatal resting state connectivity.

Distinctive cortico-striatal circuits that serve motor and cognitive functions have been recently mapped based on resting state connectivity. It has been reported that age differences in cortico-striatal connectivity relate to cognitive declines in aging. Moreover, children in their early teens (i.e., youth) already show mature motor network patterns while their cognitive networks are still developing. In the current study, we examined age differences in the frontal-striatal "cognitive" and "motor" circuits in children and adolescence, young adults (YAs), and older adults (OAs). We predicted that the strength of the "cognitive" frontal-striatal circuits would follow an inverted "U" pattern across age; children and OAs would have weaker connectivity than YAs. However, we predicted that the "motor" circuits would show less variation in connectivity strength across the lifespan. We found that most areas in both the "cognitive" and "motor" circuits showed higher connectivity in YAs than children and OAs, suggesting general inverted "U"-shaped changes across the lifespan for both the cognitive and motor frontal-striatal networks.

Association of COMT val158met and DRD2 G>T genetic polymorphisms with individual differences in motor learning and performance in female young adults.

Individuals learn new skills at different rates. Given the involvement of corticostriatal pathways in some types of learning, variations in dopaminergic transmission may contribute to these individual differences. Genetic polymorphisms of the catechol-O-methyltransferase (COMT) enzyme and dopamine receptor D2 (DRD2) genes partially determine cortical and striatal dopamine availability, respectively. Individuals who are homozygous for the COMT methionine (met) allele show reduced cortical COMT enzymatic activity, resulting in increased dopamine levels in the prefrontal cortex as opposed to individuals who are carriers of the valine (val) allele. DRD2 G-allele homozygotes benefit from a higher striatal dopamine level compared with T-allele carriers. We hypothesized that individuals who are homozygous for COMT met and DRD2 G alleles would show higher rates of motor learning. Seventy-two young healthy females (20 ± 1.9 yr) performed a sensorimotor adaptation task and a motor sequence learning task. A nonparametric mixed model ANOVA revealed that the COMT val-val group demonstrated poorer performance in the sequence learning task compared with the met-met group and showed a learning deficit in the visuomotor adaptation task compared with both met-met and val-met groups. The DRD2 TT group showed poorer performance in the sequence learning task compared with the GT group, but there was no difference between DRD2 genotype groups in adaptation rate. Although these results did not entirely come out as one might predict based on the known contribution of corticostriatal pathways to motor sequence learning, they support the role of genetic polymorphisms of COMT val158met (rs4680) and DRD2 G>T (rs 1076560) in explaining individual differences in motor performance and motor learning, dependent on task type.

Dopamine overdose hypothesis: evidence and clinical implications.

About a half a century has passed since dopamine was identified as a neurotransmitter, and it has been several decades since it was established that people with Parkinson's disease receive motor symptom relief from oral levodopa. Despite the evidence that levodopa can reduce motor symptoms, there has been a developing body of literature that dopaminergic therapy can improve cognitive functions in some patients but make them worse in others. Over the past two decades, several laboratories have shown that dopaminergic medications can impair the action of intact neural structures and impair the behaviors associated with these structures. In this review, we consider the evidence that has accumulated in the areas of reversal learning, motor sequence learning, and other cognitive tasks. The purported inverted-U shaped relationship between dopamine levels and performance is complex and includes many contributory factors. The regional striatal topography of nigrostriatal denervation is a critical factor, as supported by multimodal neuroimaging studies. A patient's individual genotype will determine the relative baseline position on this inverted-U curve. Dopaminergic pharmacotherapy and individual gene polymorphisms can affect the mesolimbic and prefrontal cortical dopaminergic functions in a comparable, inverted-U dose-response relationship. Depending on these factors, a patient can respond positively or negatively to levodopa when performing reversal learning and motor sequence learning tasks. These tasks may continue to be relevant as our society moves to increased technological demands of a digital world that requires newly learned motor sequences and adaptive behaviors to manage daily life activities.

The pattern of striatal dopaminergic denervation explains sensorimotor synchronization accuracy in Parkinson's disease.

The basal ganglia are thought to play a critical role in duration perception and production. However, experimental evidence for impaired temporal processing in Parkinson's disease (PD) patients is mixed. This study examined the association between striatal dopaminergic denervation in PD patients and sensorimotor synchronization. Twenty-eight mild-to-moderate stage PD patients synchronized finger taps to tone sequences of either 500 ms, 1000 ms or 1500 ms time intervals while ON levodopa (l-DOPA) or placebo pill (on separate test days) with the index finger of their more and less affected hands. We measured the accuracy and variability of synchronization. In a separate session, patients underwent (11)C-dihydrotetrabenazine ((11)C-DTBZ) PET scanning to measure in vivo striatal dopaminergic denervation. Patients were less accurate synchronizing to the 500 ms target time interval, compared to the 1000 ms and 1500 ms time intervals, but neither medication state nor hand affected accuracy; medication state, hand nor the target time interval affected synchronization variability. Regression analyses revealed no strong relationships between synchronization accuracy or variability and striatal dopaminergic denervation. We performed a cluster analysis on the degree of dopaminergic denervation to determine whether patient subgroup differences underlie our results. Three patient subgroups showed behavioral differences in synchronization accuracy, but not variability, paralleling their pattern of denervation. These findings provide further evidence for the role of the basal ganglia and dopamine in duration production and suggest that the degree of striatal dopaminergic denervation may explain the heterogeneity of performance between PD patients on the sensorimotor synchronization task.