A genetically defined insula-brainstem circuit selectively controls motivational vigor.

The anterior insular cortex (aIC) plays a critical role in cognitive and motivational control of behavior, but the underlying neural mechanism remains elusive. Here, we show that aIC neurons expressing Fezf2 (aICFezf2), which are the pyramidal tract neurons, signal motivational vigor and invigorate need-seeking behavior through projections to the brainstem nucleus tractus solitarii (NTS). aICFezf2 neurons and their postsynaptic NTS neurons acquire anticipatory activity through learning, which encodes the perceived value and the vigor of actions to pursue homeostatic needs. Correspondingly, aIC → NTS circuit activity controls vigor, effort, and striatal dopamine release but only if the action is learned and the outcome is needed. Notably, aICFezf2 neurons do not represent taste or valence. Moreover, aIC → NTS activity neither drives reinforcement nor influences total consumption. These results pinpoint specific functions of aIC → NTS circuit for selectively controlling motivational vigor and suggest that motivation is subserved, in part, by aIC's top-down regulation of dopamine signaling.

Beta and theta oscillations track effort and previous reward in the human basal ganglia and prefrontal cortex during decision making.

Choosing whether to exert effort to obtain rewards is fundamental to human motivated behavior. However, the neural dynamics underlying the evaluation of reward and effort in humans is poorly understood. Here, we report an exploratory investigation into this with chronic intracranial recordings from the prefrontal cortex (PFC) and basal ganglia (BG; subthalamic nuclei and globus pallidus) in people with Parkinson's disease performing a decision-making task with offers that varied in levels of reward and physical effort required. This revealed dissociable neural signatures of reward and effort, with BG beta (12 to 20 Hz) oscillations tracking effort on a single-trial basis and PFC theta (4 to 7 Hz) signaling previous trial reward, with no effects of net subjective value. Stimulation of PFC increased overall acceptance of offers and sensitivity to reward while decreasing the impact of effort on choices. This work uncovers oscillatory mechanisms that guide fundamental decisions to exert effort for reward across BG and PFC, supports a causal role of PFC for such choices, and seeds hypotheses for future studies.

Global expansion of marine protected areas and the redistribution of fishing effort.

The expansion of marine protected areas (MPAs) is a core focus of global conservation efforts, with the "30x30" initiative to protect 30% of the ocean by 2030 serving as a prominent example of this trend. We consider a series of proposed MPA network expansions of various sizes, and we forecast the impact this increase in protection would have on global patterns of fishing effort. We do so by building a predictive machine learning model trained on a global dataset of satellite-based fishing vessel monitoring data, current MPA locations, and spatiotemporal environmental, geographic, political, and economic features. We then use this model to predict future fishing effort under various MPA expansion scenarios compared to a business-as-usual counterfactual scenario that includes no new MPAs. The difference between these scenarios represents the predicted change in fishing effort associated with MPA expansion. We find that regardless of the MPA network objectives or size, fishing effort would decrease inside the MPAs, though by much less than 100%. Moreover, we find that the reduction in fishing effort inside MPAs does not simply redistribute outside-rather, fishing effort outside MPAs would also decline. The overall magnitude of the predicted decrease in global fishing effort principally depends on where networks are placed in relation to existing fishing effort. MPA expansion will lead to a global redistribution of fishing effort that should be accounted for in network design, implementation, and impact evaluation.

Effort Foraging Task reveals positive correlation between individual differences in the cost of cognitive and physical effort in humans.

Effort-based decisions, in which people weigh potential future rewards against effort costs required to achieve those rewards involve both cognitive and physical effort, though the mechanistic relationship between them is not yet understood. Here, we use an individual differences approach to isolate and measure the computational processes underlying effort-based decisions and test the association between cognitive and physical domains. Patch foraging is an ecologically valid reward rate maximization problem with well-developed theoretical tools. We developed the Effort Foraging Task, which embedded cognitive or physical effort into patch foraging, to quantify the cost of both cognitive and physical effort indirectly, by their effects on foraging choices. Participants chose between harvesting a depleting patch, or traveling to a new patch that was costly in time and effort. Participants' exit thresholds (reflecting the reward they expected to receive by harvesting when they chose to travel to a new patch) were sensitive to cognitive and physical effort demands, allowing us to quantify the perceived effort cost in monetary terms. The indirect sequential choice style revealed effort-seeking behavior in a minority of participants (preferring high over low effort) that has apparently been missed by many previous approaches. Individual differences in cognitive and physical effort costs were positively correlated, suggesting that these are perceived and processed in common. We used canonical correlation analysis to probe the relationship of task measures to self-reported affect and motivation, and found correlations of cognitive effort with anxiety, cognitive function, behavioral activation, and self-efficacy, but no similar correlations with physical effort.

Generalized Encoding of the Relative Subjective Value of Cognitive Effort in the Dorsal ACC.

Making choices about whether and when to engage cognitive effort are a common feature of everyday experience, with important consequences for academic, career, and health outcomes. Yet, despite their hypothesized importance, very little is understood about the underlying mechanisms that support this form of human cost-benefit decision-making. To investigate these mechanisms, we used the Cognitive Effort Discounting Paradigm (Cog-ED) during fMRI scanning to precisely quantify the neural encoding of varying cognitive effort demands relative to reward outcomes, within two distinct cognitive domains (working memory, speech comprehension). The findings provide strong evidence that the dorsal anterior cingulate cortex (dACC) plays a central and selective role in this decision-making process. Trial-by-trial modulations in dACC activation tracked the relative subjective value of the low-effort, low-reward option, with the strongest activity occurring when this was of greater value than the high-effort, high-reward option. In contrast, dACC activity was not modulated by decision difficulty, though such effects were found in other frontoparietal regions. Critically, dACC activity was also strongly correlated across the two decision-making task domains and further predicted subsequent choice behavior in both. Together, the results suggest that dACC activity modulation reflects a domain-general valuation comparison mechanism, which acts to bias participants away from decisions to engage in cognitive effort, when the perceived subjective costs of such engagement outweigh the reward-related benefits. These findings complement work in other cost domains and species by pointing to a clear role of the dACC in representing subjective value differences between choice options during cost-benefit decision-making.

Slowing of Movements in Healthy Aging as a Rational Economic Response to an Elevated Effort Landscape.

Why do we move slower as we grow older? The reward circuits of the brain, which tend to invigorate movements, decline with aging, raising the possibility that reduced vigor is due to the diminishing value that our brain assigns to movements. However, as we grow older, it also becomes more effortful to make movements. Is age-related slowing principally a consequence of increased effort costs from the muscles, or reduced valuation of reward by the brain? Here, we first quantified the cost of reaching via metabolic energy expenditure in human participants (male and female), and found that older adults consumed more energy than the young at a given speed. Thus, movements are objectively more costly for older adults. Next, we observed that when reward increased, older adults, like the young, responded by initiating their movements earlier. Yet, unlike the young, they were unwilling to increase their movement speed. Was their reluctance to reach quicker for rewards due to the increased effort costs, or because they ascribed less value to the movement? Motivated by a mathematical model, we next made the young experience a component of aging by making their movements more effortful. Now the young responded to reward by reacting faster but chose not to increase their movement speed. This suggests that slower movements in older adults are partly driven by an adaptive response to an elevated effort landscape. Moving slower may be a rational economic response the brain is making to mitigate the elevated effort costs that accompany aging.

Trying Harder: How Cognitive Effort Sculpts Neural Representations during Working Memory.

While the exertion of mental effort improves performance on cognitive tasks, the neural mechanisms by which motivational factors impact cognition remain unknown. Here, we used fMRI to test how changes in cognitive effort, induced by changes in task difficulty, impact neural representations of working memory (WM). Participants (both sexes) were precued whether WM difficulty would be hard or easy. We hypothesized that hard trials demanded more effort as a later decision required finer mnemonic precision. Behaviorally, pupil size was larger and response times were slower on hard compared with easy trials suggesting our manipulation of effort succeeded. Neurally, we observed robust persistent activity during delay periods in the prefrontal cortex (PFC), especially during hard trials. Yet, details of the memoranda could not be decoded from patterns in prefrontal activity. In the patterns of activity in the visual cortex, however, we found strong decoding of memorized targets, where accuracy was higher on hard trials. To potentially link these across-region effects, we hypothesized that effort, carried by persistent activity in the PFC, impacts the quality of WM representations encoded in the visual cortex. Indeed, we found that the amplitude of delay period activity in the frontal cortex predicted decoded accuracy in the visual cortex on a trial-wise basis. These results indicate that effort-related feedback signals sculpt population activity in the visual cortex, improving mnemonic fidelity.

Value Estimation versus Effort Mobilization: A General Dissociation between Ventromedial and Dorsomedial Prefrontal Cortex.

Deciding on a course of action requires both an accurate estimation of option values and the right amount of effort invested in deliberation to reach sufficient confidence in the final choice. In a previous study, we have provided evidence, across a series of judgment and choice tasks, for a dissociation between the ventromedial prefrontal cortex (vmPFC), which would represent option values, and the dorsomedial prefrontal cortex (dmPFC), which would represent the duration of deliberation. Here, we first replicate this dissociation and extend it to the case of an instrumental learning task, in which 24 human volunteers (13 women) choose between options associated with probabilistic gains and losses. According to fMRI data recorded during decision-making, vmPFC activity reflects the sum of option values generated by a reinforcement learning model and dmPFC activity the deliberation time. To further generalize the role of the dmPFC in mobilizing effort, we then analyze fMRI data recorded in the same participants while they prepare to perform motor and cognitive tasks (squeezing a handgrip or making numerical comparisons) to maximize gains or minimize losses. In both cases, dmPFC activity is associated with the output of an effort regulation model, and not with response time. Taken together, these results strengthen a general theory of behavioral control that implicates the vmPFC in the estimation of option values and the dmPFC in the energization of relevant motor and cognitive processes.

The effect of Huntington's disease on cognitive and physical motivation.

Apathy is one of the most common neuropsychiatric features of Huntington's disease. A hallmark of apathy is diminished goal-directed behaviour, which is characterized by a lower motivation to engage in cognitively or physically effortful actions. However, it remains unclear whether this reduction in goal-directed behaviour is driven primarily by a motivational deficit and/or is secondary to the progressive cognitive and physical deficits that accompany more advanced disease. We addressed this question by testing 17 individuals with manifest Huntington's disease and 22 age-matched controls on an effort-based decision-making paradigm. Participants were first trained on separate cognitively and physically effortful tasks and provided explicit feedback about their performance. Next, they chose on separate trials how much effort they were willing to exert in each domain in return for varying reward. At the conclusion of the experiment, participants were asked to rate their subjective perception of task load. In the cognitive task, the Huntington's disease group were more averse to cognitive effort than controls. Although the Huntington's disease group were more impaired than controls on the task itself, their greater aversion to cognitive effort persisted even after controlling for task performance. This suggests that the lower levels of cognitive motivation in the Huntington's disease group relative to controls was most likely driven by a primary motivational deficit. In contrast, both groups expressed a similar preference for physical effort. Importantly, the similar levels of physical motivation across both groups occurred even though participants with Huntington's disease performed objectively worse than controls on the physical effort task, and were aware of their performance through explicit feedback on each trial. This indicates that the seemingly preserved level of physical motivation in Huntington's disease was driven by a willingness to engage in physically effortful actions despite a reduced capacity to do so. Finally, the Huntington's disease group provided higher ratings of subjective task demand than controls for the cognitive (but not physical) effort task and when assessing the mental (but not the physical) load of each task. Together, these results revealed a dissociation in cognitive and physical motivation deficits between Huntington's disease and controls, which were accompanied by differences in how effort was subjectively perceived by the two groups. This highlights that motivation is the final manifestation of a complex set of mechanisms involved in effort processing, which are separable across different domains of behaviour. These findings have important clinical implications for the day-to-day management of apathy in Huntington's disease.

Decision cost hypersensitivity underlies Huntington's disease apathy.

The neuropsychiatric syndrome of apathy is now recognized to be a common and disabling condition in Huntington's disease (HD). However, the mechanisms underlying it are poorly understood. One way to investigate apathy is to utilise a theoretical framework of normal motivated behaviour, to determine where breakdown has occurred in people with this behavioural disruption. A fundamental computation underlying motivated, goal-directed behaviour across species is weighing up the costs and rewards associated with actions. Here, we asked whether people with apathy are more sensitive to costs of actions (physical effort and time delay), less sensitive to rewarding outcomes, or both. Based on the unique anatomical substrates associated with HD pathology, we hypothesised that a general hypersensitivity to costs would underpin HD apathy. Genetically confirmed carriers of the expanded Huntingtin gene (premanifest to mild motor manifest disease (n=53) were compared to healthy controls (n = 38). Participants performed a physical effort-based decision-making task (Apple Gathering Task) and a delay discounting task (Money Choice Questionnaire). Choice data was analysed using linear regression and drift diffusion models that also accounted for the time taken to make decisions. Apathetic people with HD accepted fewer offers overall on the Apple Gathering Task, specifically driven by increased sensitivity to physical effort costs, and not explained by motor severity, mood, cognition, or medication. Drift diffusion modelling provided further evidence of effort hypersensitivity, with apathy associated with a faster drift rate towards rejecting offers as a function of varying effort. Increased delay sensitivity was also associated with apathy, both when analysing raw choice and also drift rate, where there was moderate evidence of HD apathy drifting faster towards the immediately available (low cost) option. Furthermore, the effort and delay sensitivity parameters from these tasks were positively correlated. The results demonstrate a clear mechanism for apathy in HD, cost hypersensitivity, which manifests in both the effort and time costs associated with actions towards rewarding goals. This suggests that HD pathology may cause a domain-general disruption of cost processing, which is distinct to apathy occurrence in other brain disorders, and may require different therapeutic approaches.

Aberrant effort-based reward dynamics in anhedonia.

Anhedonia is a transdiagnostic symptom and associated with a spectrum of reward deficits among which the motivational dysfunction is poorly understood. Previous studies have established the abnormal cost-benefit trade-off as a contributor to motivational deficits in anhedonia and its relevant psychiatric diseases. However, it remains elusive how the anhedonic neural dynamics underlying reward processing are modulated by effort expenditure. Using an effort-based monetary incentive delay task, the current event-related potential study examined the neural dynamics underlying the effort-reward interplay in anhedonia using a nonclinical sample who scored high or low on an anhedonia questionnaire. We found that effort prospectively decreased reward effect on the contingent variation negativity and the target-P3 but retrospectively enhanced outcome effect on the feedback-P3 following effort expenditure. Compared to the low-anhedonia group, the high-anhedonia group displayed a diminished effort effect on the target-P3 during effort expenditure and an increased effort-enhancement effect for neutral trials during the feedback-P3 period following effort expenditure. Our findings suggest that anhedonia is associated with an inefficient control and motivation allocation along the efforted-based reward dynamics from effort preparation to effort production.

Effort expenditure modulates feedback evaluations involving self-other agreement: evidence from brain potentials and neural oscillations.

The influence of effort expenditure on the subjective value in feedback involving material reward has been the focus of previous research. However, little is known about the impact of effort expenditure on subjective value evaluations when feedback involves reward that is produced in the context of social interaction (e.g. self-other agreement). Moreover, how effort expenditure influences confidence (second-order subjective value) in feedback evaluations remains unclear. Using electroencephalography, this study aimed to address these questions. Event-related potentials showed that, after exerting high effort, participants exhibited increased reward positivity difference in response to self-other (dis)agreement feedback. After exerting low effort, participants reported high confidence, and the self-other disagreement feedback evoked a larger P3a. Time-frequency analysis showed that the high-effort task evoked increased frontal midline theta power. In the low (vs. high)-effort task, the frontal midline delta power for self-other disagreement feedback was enhanced. These findings suggest that, at the early feedback evaluation stage, after exerting high effort, individuals exhibit an increased sensitivity of subjective value evaluation in response to self-other agreement feedback. At the later feedback evaluation stage, after completing the low-effort task, the self-other disagreement feedback violates the individuals'high confidence and leads to a metacognitive mismatch.

Rewarding cognitive effort increases the intrinsic value of mental labor.

Current models of mental effort in psychology, behavioral economics, and cognitive neuroscience typically suggest that exerting cognitive effort is aversive, and people avoid it whenever possible. The aim of this research was to challenge this view and show that people can learn to value and seek effort intrinsically. Our experiments tested the hypothesis that effort-contingent reward in a working-memory task will induce a preference for more demanding math tasks in a transfer phase, even though participants were aware that they would no longer receive any reward for task performance. In laboratory Experiment 1 (n = 121), we made reward directly contingent on mobilized cognitive effort as assessed via cardiovascular measures (ß-adrenergic sympathetic activity) during the training task. Experiments 2a to 2e (n = 1,457) were conducted online to examine whether the effects of effort-contingent reward on subsequent demand seeking replicate and generalize to community samples. Taken together, the studies yielded reliable evidence that effort-contingent reward increased participants' demand seeking and preference for the exertion of cognitive effort on the transfer task. Our findings provide evidence that people can learn to assign positive value to mental effort. The results challenge currently dominant theories of mental effort and provide evidence and an explanation for the positive effects of environments appreciating effort and individual growth on people's evaluation of effort and their willingness to mobilize effort and approach challenging tasks.

Amphetamines Improve the Motivation to Invest Effort in Attention-Deficit/Hyperactivity Disorder.

Prevailing frameworks propose that a key feature of attention-deficit/hyperactivity disorder (ADHD) is lower motivation. An important component of motivation is the willingness to engage in cognitively or physically effortful behavior. However, the degree to which effort sensitivity is impaired in ADHD has rarely been tested, and the efficacy of stimulant medication in ameliorating any such impairments is unclear. Here, we tested 20 individuals with ADHD (11 males, 9 females) who were managed with amphetamine-based medication (dexamfetamine, lisdexamfetamine), and 24 controls (8 males, 16 females). Individuals with ADHD were tested over two counterbalanced sessions, ON and OFF their usual amphetamine-based medication. In each session, participants performed an effort-based decision-making task, in which they were required to choose how much cognitive or physical effort they were willing to engage in return for reward. Our results revealed three main findings. First, individuals with ADHD had lower motivation relative to controls to invest effort in both the cognitive and physical domains. Second, amphetamine increased motivation uniformly across both domains. Finally, the net effect of amphetamine treatment was to mostly restore motivation across both domains of effort relative to healthy controls. These data provide clear evidence for a heightened sensitivity to both cognitive and physical effort in ADHD, and reveal the efficacy of amphetamine-based drugs in restoring effort sensitivity to levels similar to controls. These findings confirm the existence of reduced motivational drive in ADHD, and more broadly provide direct causal evidence for a domain-general role of catecholamines in motivating effortful behavior.SIGNIFICANCE STATEMENT A core feature of attention-deficit/hyperactivity disorder (ADHD) is thought to be a heightened aversion to effort. Surprisingly, however, the degree to which effort sensitivity is impaired in ADHD has rarely been tested. More broadly, the relative efficacy of catecholamines in motivating the investment of cognitive and physical effort is unclear. We tested 20 individuals with ADHD ON and OFF amphetamines, and compared their behavior on an effort-based decision-making task to 24 controls. When tested OFF medication, the ADHD group was less cognitively and physically motivated than controls. However, amphetamines led to a comparable increase in motivation across both domains. This demonstrates the efficacy of catecholamines in facilitating domain-general effort, and highlights the broader potential of such drugs to treat disorders of motivation.

Boosting Serotonin Increases Information Gathering by Reducing Subjective Cognitive Costs.

Serotonin is implicated in the valuation of aversive costs, such as delay or physical effort. However, its role in governing sensitivity to cognitive effort, for example, deliberation costs during information gathering, is unclear. We show that treatment with a serotonergic antidepressant in healthy human individuals of either sex enhances a willingness to gather information when trying to maximize reward. Using computational modeling, we show this arises from a diminished sensitivity to subjective deliberation costs during the sampling process. This result is consistent with the notion that serotonin alleviates sensitivity to aversive costs in a domain-general fashion, with implications for its potential contribution to a positive impact on motivational deficits in psychiatric disorders.SIGNIFICANCE STATEMENT Gathering information about the world is essential for successfully navigating it. However, sampling information is costly, and we need to balance between gathering too little and too much information. The neurocomputational mechanisms underlying this arbitration between a putative gain, such as reward, and the associated costs, such as allocation of cognitive resources, remain unclear. In this study, we show that week-long daily treatment with a serotonergic antidepressant enhances a willingness to gather information when trying to maximize reward. Computational modeling indicates this arises from a reduced perception of aversive costs, rendering information gathering less cognitively effortful. This finding points to a candidate mechanism by which serotonergic treatment might help alleviate motivational deficits in a range of mental illnesses.

Pupil Dilation and Microsaccades Provide Complementary Insights into the Dynamics of Arousal and Instantaneous Attention during Effortful Listening.

Listening in noisy environments requires effort- the active engagement of attention and other cognitive abilities- as well as increased arousal. The ability to separately quantify the contribution of these components is key to understanding the dynamics of effort and how it may change across listening situations and in certain populations. We concurrently measured two types of ocular data in young participants (both sexes): pupil dilation (PD; thought to index arousal aspects of effort) and microsaccades (MS; hypothesized to reflect automatic visual exploratory sampling), while they performed a speech-in-noise task under high- (HL) and low- (LL) listening load conditions. Sentences were manipulated so that the behaviorally relevant information (keywords) appeared at the end (Experiment 1) or beginning (Experiment 2) of the sentence, resulting in different temporal demands on focused attention. In line with previous reports, PD effects were associated with increased dilation under load. We observed a sustained difference between HL and LL conditions, consistent with increased phasic and tonic arousal. Importantly we show that MS rate was also modulated by listening load. This was manifested as a reduced MS rate in HL relative to LL. Critically, in contrast to the sustained difference seen for PD, MS effects were localized in time, specifically during periods when demands on auditory attention were greatest. These results demonstrate that auditory selective attention interfaces with the mechanisms controlling MS generation, establishing MS as an informative measure, complementary to PD, with which to quantify the temporal dynamics of auditory attentional processing under effortful listening conditions.SIGNIFICANCE STATEMENT Listening effort, reflecting the "cognitive bandwidth" deployed to effectively process sound in adverse environments, contributes critically to listening success. Understanding listening effort and the processes involved in its allocation is a major challenge in auditory neuroscience. Here, we demonstrate that microsaccade rate can be used to index a specific subcomponent of listening effort, the allocation of instantaneous auditory attention, that is distinct from the modulation of arousal indexed by pupil dilation (currently the dominant measure of listening effort). These results reveal the push-pull process through which auditory attention interfaces with the (visual) attention network that controls microsaccades, establishing microsaccades as a powerful tool for measuring auditory attention and its deficits.

Eye Movements Decrease during Effortful Speech Listening.

Hearing impairment affects many older adults but is often diagnosed decades after speech comprehension in noisy situations has become effortful. Accurate assessment of listening effort may thus help diagnose hearing impairment earlier. However, pupillometry-the most used approach to assess listening effort-has limitations that hinder its use in practice. The current study explores a novel way to assess listening effort through eye movements. Building on cognitive and neurophysiological work, we examine the hypothesis that eye movements decrease when speech listening becomes challenging. In three experiments with human participants from both sexes, we demonstrate, consistent with this hypothesis, that fixation duration increases and spatial gaze dispersion decreases with increasing speech masking. Eye movements decreased during effortful speech listening for different visual scenes (free viewing, object tracking) and speech materials (simple sentences, naturalistic stories). In contrast, pupillometry was less sensitive to speech masking during story listening, suggesting pupillometric measures may not be as effective for the assessments of listening effort in naturalistic speech-listening paradigms. Our results reveal a critical link between eye movements and cognitive load, suggesting that neural activity in the brain regions that support the regulation of eye movements, such as frontal eye field and superior colliculus, are modulated when listening is effortful.SIGNIFICANCE STATEMENT Assessment of listening effort is critical for early diagnosis of age-related hearing loss. Pupillometry is most used but has several disadvantages. The current study explores a novel way to assess listening effort through eye movements. We examine the hypothesis that eye movements decrease when speech listening becomes effortful. We demonstrate, consistent with this hypothesis, that fixation duration increases and gaze dispersion decreases with increasing speech masking. Eye movements decreased during effortful speech listening for different visual scenes (free viewing, object tracking) and speech materials (sentences, naturalistic stories). Our results reveal a critical link between eye movements and cognitive load, suggesting that neural activity in brain regions that support the regulation of eye movements are modulated when listening is effortful.

From Motivation to Action: Action Cost Better Predicts Changes in Premovement Beta-Band Activity than Speed.

Although premovement beta-band event-related desynchronization (ß-ERD; 13-30 Hz) from sensorimotor regions is modulated by movement speed, current evidence does not support a strict monotonic association between the two. Given that ß-ERD is thought to increase information encoding capacity, we tested the hypothesis that it might be related to the expected neurocomputational cost of movement, here referred to as action cost. Critically, action cost is greater both for slow and fast movements compared with a medium or "preferred" speed. Thirty-one right-handed participants performed a speed-controlled reaching task while recording their EEG. Results revealed potent modulations of beta power as a function of speed, with ß-ERD being significantly greater both for movements performed at high and low speeds compared with medium speed. Interestingly, medium-speed movements were more often chosen by participants than low-speed and high-speed movements, suggesting that they were evaluated as less costly. In line with this, modeling of action cost revealed a pattern of modulation across speed conditions that strikingly resembled the one found for ß-ERD. Indeed, linear mixed models showed that estimated action cost predicted variations of ß-ERD significantly better than speed. This relationship with action cost was specific to beta power, as it was not found when averaging activity in the mu band (8-12 Hz) and gamma band (31-49 Hz) bands. These results demonstrate that increasing ß-ERD may not merely speed up movements, but instead facilitate the preparation of high-speed and low-speed movements through the allocation of additional neural resources, thereby enabling flexible motor control.SIGNIFICANCE STATEMENT Heightened beta activity has been associated with movement slowing in Parkinson's disease, and modulations of beta activity are commonly used to decode movement parameters in brain-computer interfaces. Here we show that premovement beta activity is better explained by the neurocomputational cost of the action rather than its speed. Instead of being interpreted as a mere reflection of changes in movement speed, premovement changes in beta activity might therefore be used to infer the amount of neural resources that are allocated for motor planning.

Effects of reward and effort history on decision making and movement vigor during foraging.

During foraging, animals explore a site and harvest reward and then abandon that site and travel to the next opportunity. One aspect of this behavior involves decision making, and the other involves movement control. These two aspects of behavior may be linked via an underlying desire to maximize a single normative utility: the sum of all rewards acquired, minus all efforts expended, divided by time. According to this theory, the history of rewards, and not just its immediate availability, should dictate how long one should stay and harvest reward and how vigorously one should travel to the next opportunity. We tested this theory in a series of experiments in which humans used their hand to harvest tokens at a reward patch and then used their arm to reach toward another patch. After a history of high rewards, the subjects not only shortened their harvest duration but also moved more vigorously toward the next reward opportunity. In contrast, after a history of high effort they lengthened their harvest duration but reduced their movement vigor, reaching more slowly to the next reward site. Thus, a history of high reward or low effort biased decisions by promoting early abandonment of the reward site and biased movements by promoting vigor.NEW & NOTEWORTHY Much of life is spent foraging. Whereas previous work has focused on the decision regarding time spent harvesting from a reward patch, here we test the idea that both decision making and movement control are tuned to optimize the net rate of reward in an environment. Our results show that movement patterns reflect not just immediate expectations but also past experiences in the environment, providing fundamental insight into the factors governing volitional control of arm movements.

Human foot force informs balance control strategies when standing on a narrow beam.

Despite the abundance of studies on the control of standing balance, insights about the roles of biomechanics and neural control have been limited. Previous work introduced an analysis combining the direction and orientation of foot-ground forces. The "intersection point" of the lines of actions of these forces exhibited a consistent pattern across healthy, young subjects when computed for different frequency components of the center of pressure signal. To investigate the control strategy of quiet stance, we applied this intersection point analysis to experimental data of 15 healthy, young subjects balancing in tandem stance on a narrow beam and on the ground. Data from the sagittal and frontal planes were analyzed separately. The task was modeled as a double-inverted pendulum controlled by an optimal controller with torque-actuated ankle and hip joints and additive white noise. To test our prediction that the controller that minimized overall joint effort would yield the best fit across the tested conditions and planes of analyses, experimental results were compared with simulation outcomes. The controller that minimized overall effort produced the best fit in both balance conditions and planes of analyses. For some conditions, the relative penalty on the hip and ankle joints varied in a way relevant to the balance condition or to the plane of analysis. These results suggest that unimpaired quiet balance in a challenging environment can be best described by a controller that maintains minimal effort through the adjustment of relative ankle and hip joint torques. NEW & NOTEWORTHY This study explored balance control in humans during a challenging task using the novel intersection point analysis, based on foot-ground force direction and point of application. Experimental data of subjects standing on a narrow beam in tandem stance were compared with modeling results of a double-inverted pendulum. The analysis showed that individuals minimized effort by adjusting ankle and hip torques, shedding light on the interplay of biomechanics and neural control in maintaining balance.