Visuomotor Control Task Research Articles

Advanced Force Coordination of Lower Extremities During Visuomotor Control Task in Soccer Players

ABSTRACT Purpose: This study is aimed to determine specific bilateral lower extremity motor performances and coordination patterns in soccer players with healthy controls using the bilateral force control paradigm. We hypothesized that soccer players would show more advanced bilateral force control performances than untrained controls. Methods: Participants were 13 university soccer players and 13 healthy controls. Each group performed bilateral ankle dorsiflexion force control tasks across two vision conditions (i.e., vision and no-vision) and two targeted force levels (i.e., 10% and 40% maximum voluntary contraction). We calculated force accuracy, variability, and symmetry to assess force control performances. To estimate bilateral force coordination, we calculated Pearson’s correlation coefficients between feet-within a single trial and considered uncontrolled manifold variables across multiple trials. In the no-vision condition, we performed secondary analyses for initial force control patters after removal of visual feedback. Results: There were no significant group differences in bilateral force accuracy and variability but, compared to the control group, soccer players showed higher force symmetry between left and right ankle dorsiflexion forces. For force coordination between feet, soccer players revealed more negative values of the correlation coefficient and greater good variability from the uncontrolled manifold analysis than those for the control group. The secondary analysis revealed no significant group differences in the time until force drift and amount of force adjustments. Conclusions: Soccer players have more compensatory and flexible interlimb force coordination strategies between feet.

Eye movements during motor imagery and execution reveal different visuomotor control strategies in manual interception.

Previous research has investigated the degree of congruency in gaze metrics between action execution (AE) and motor imagery (MI) for similar manual tasks. Although eye movement dynamics seem to be limited to relatively simple actions toward static objects, there is little evidence of how gaze parameters change during imagery as a function of more dynamic spatial and temporal task demands. This study examined the similarities and differences in eye movements during AE and MI for an interception task. Twenty-four students were asked to either mentally simulate or physically intercept a moving target on a computer display. Smooth pursuit, saccades, and response time were compared between the two conditions. The results show that MI was characterized by higher smooth pursuit gain and duration while no meaningful differences were found in the other parameters. The findings indicate that eye movements during imagery are not simply a duplicate of what happens during actual performance. Instead, eye movements appear to vary as a function of the interaction between visuomotor control strategies and task demands.

Examining the role of attentional allocation in working memory precision with pupillometry in children and adults

Working memory (WM) precision, or the fidelity with which items can be remembered, is an important aspect of WM capacity that increases over childhood. Why individuals are more or less precise from moment to moment and why WM becomes more stable with age are not yet fully understood. Here, we examined the role of attentional allocation in visual WM precision in children aged 8 to 13 years and young adults aged 18 to 27 years, as measured by fluctuations in pupil dilation during stimulus encoding and maintenance. Using mixed models, we examined intraindividual links between change in pupil diameter and WM precision across trials and the role of developmental differences in these associations. Through probabilistic modeling of error distributions and the inclusion of a visuomotor control task, we isolated mnemonic precision from other cognitive processes. We found an age-related increase in mnemonic precision that was independent of guessing behavior, serial position effects, fatigue or loss of motivation across the experiment, and visuomotor processes. Trial-by-trial analyses showed that trials with smaller changes in pupil diameter during encoding and maintenance predicted more precise responses than trials with larger changes in pupil diameter within individuals. At encoding, this relationship was stronger for older participants. Furthermore, the pupil–performance coupling grew across the delay period—particularly or exclusively for adults. These results suggest a functional link between pupil fluctuations and WM precision that grows over development; visual details may be stored more faithfully when attention is allocated efficiently to a sequence of objects at encoding and throughout a delay period.

Transcranial direct current stimulation over the posterior parietal cortex improves visuomotor performance and proprioception in the lower extremities

The purpose of this study was to examine whether anodal transcranial direct current stimulation (a-tDCS) over the posterior parietal cortex (PPC) could affect visuomotor performance and proprioception in the lower extremities. We evaluated visuomotor performance in 15 healthy volunteers using a visuomotor control task by plantar dorsiflexion of the ankle joint, and calculated the absolute difference between the target and measured angle. In addition, we evaluated proprioception using a joint position matching task. During the task, the subject reproduced the ankle joint plantar dorsiflexion angle presented by the examiner. We calculated the absolute difference between the presented and measured angles (absolute error) and the variation of measured angles (variable error). Simultaneously, a-tDCS (1.5 mA, 15 min) or sham stimulation was applied to the right PPC. We observed that the absolute error of the visuomotor control task and the variable error of the joint position matching task significantly decreased after a-tDCS. However, the absolute error of the joint position matching task was not affected. This study suggests that a-tDCS over the PPC improves visuomotor performance and reduces the variable error in the joint position matching task.

Intuitive physical reasoning about objects' masses transfers to a visuomotor decision task consistent with Newtonian physics.

While interacting with objects during every-day activities, e.g. when sliding a glass on a counter top, people obtain constant feedback whether they are acting in accordance with physical laws. However, classical research on intuitive physics has revealed that people's judgements systematically deviate from predictions of Newtonian physics. Recent research has explained at least some of these deviations not as consequence of misconceptions about physics but instead as the consequence of the probabilistic interaction between inevitable perceptual uncertainties and prior beliefs. How intuitive physical reasoning relates to visuomotor actions is much less known. Here, we present an experiment in which participants had to slide pucks under the influence of naturalistic friction in a simulated virtual environment. The puck was controlled by the duration of a button press, which needed to be scaled linearly with the puck's mass and with the square-root of initial distance to reach a target. Over four phases of the experiment, uncertainties were manipulated by altering the availability of sensory feedback and providing different degrees of knowledge about the physical properties of pucks. A hierarchical Bayesian model of the visuomotor interaction task incorporating perceptual uncertainty and press-time variability found substantial evidence that subjects adjusted their button-presses so that the sliding was in accordance with Newtonian physics. After observing collisions between pucks, which were analyzed with a hierarchical Bayesian model of the perceptual observation task, subjects transferred the relative masses inferred perceptually to adjust subsequent sliding actions. Crucial in the modeling was the inclusion of a cost function, which quantitatively captures participants' implicit sensitivity to errors due to their motor variability. Taken together, in the present experiment we find evidence that our participants transferred their intuitive physical reasoning to a subsequent visuomotor control task consistent with Newtonian physics and weighed potential outcomes with a cost functions based on their knowledge about their own variability.

Effects on motor learning of transcranial alternating current stimulation applied over the primary motor cortex and cerebellar hemisphere

Transcranial alternating current stimulation (tACS) is a non-invasive method of brain stimulation that modulates oscillatory neural activity in the cortical area under the electrodes. Gamma (γ)-tACS applied over the primary motor cortex (M1) and cerebellar hemisphere is known to improve motor performance; however, it is not yet known whether it affects motor learning. Thus, here we investigated whether γ-tACS applied over the M1 and cerebellar hemisphere affects motor learning. This study involved 30 healthy subjects (14 females, 16 males) performing a visuomotor control task (eight trials) during an administration of either γ-tACS or a sham stimulation (15 subjects per condition) over their right M1 and left cerebellar hemisphere. Each subject performed five trials after 24 h. The motor learning efficiency, motor learning retention and re-motor learning efficiency in each condition were compared. The motor learning retention in the γ-tACS condition was significantly higher than that in the sham condition (p = 0.031). Thus, subjects who were administered γ-tACS maintained their motor performance the next day better than sham-stimulated subjects. There was no significant difference between the conditions in the motor learning efficiency and those in the re-motor learning efficiency. Our results demonstrate that γ-tACS administered over the M1 and cerebellar hemisphere during a motor learning task can enhance motor learning retention.

Wireless Sensing of Lower Lip and Thumb-Index Finger 'Ramp-and-Hold' Isometric Force Dynamics in a Small Cohort of Unilateral MCA Stroke: Discussion of Preliminary Findings.

Automated wireless sensing of force dynamics during a visuomotor control task was used to rapidly assess residual motor function during finger pinch (right and left hand) and lower lip compression in a cohort of seven adult males with chronic, unilateral middle cerebral artery (MCA) stroke with infarct confirmed by anatomic magnetic resonance imaging (MRI). A matched cohort of 25 neurotypical adult males served as controls. Dependent variables were extracted from digitized records of ‘ramp-and-hold’ isometric contractions to target levels (0.25, 0.5, 1, and 2 Newtons) presented in a randomized block design; and included force reaction time, peak force, and dF/dtmax associated with force recruitment, and end-point accuracy and variability metrics during the contraction hold-phase (mean, SD, criterion percentage ‘on-target’). Maximum voluntary contraction force (MVCF) was also assessed to establish the force operating range. Results based on linear mixed modeling (LMM, adjusted for age and handedness) revealed significant patterns of dissolution in fine force regulation among MCA stroke participants, especially for the contralesional thumb-index finger followed by the ipsilesional digits, and the lower lip. For example, the contralesional thumb-index finger manifest increased reaction time, and greater overshoot in peak force during recruitment compared to controls. Impaired force regulation among MCA stroke participants during the contraction hold-phase was associated with significant increases in force SD, and dramatic reduction in the ability to regulate force output within prescribed target force window (±5% of target). Impaired force regulation during contraction hold-phase was greatest in the contralesional hand muscle group, followed by significant dissolution in ipsilateral digits, with smaller effects found for lower lip. These changes in fine force dynamics were accompanied by large reductions in the MVCF with the LMM marginal means for contralesional and ipsilesional pinch forces at just 34.77% (15.93 N vs. 45.82 N) and 66.45% (27.23 N vs. 40.98 N) of control performance, respectively. Biomechanical measures of fine force and MVCF performance in adult stroke survivors provide valuable information on the profile of residual motor function which can help inform clinical treatment strategies and quantitatively monitor the efficacy of rehabilitation or neuroprotection strategies.

The effect of gamma tACS over the M1 region and cerebellar hemisphere does not depend on current intensity

Transcranial alternating current stimulation (tACS) has been shown to modulate neural connectivity in the cortical area under experimental electrodes. Although gamma tACS over the M1 region and the cerebellar hemisphere has been shown to improve motor performance, the optimal stimulation method has not been clarified. In this study, we aimed to clarify whether the effect of gamma tACS over M1 and the cerebellar hemisphere depends on current intensity. Twenty healthy adults performed a visuomotor control task using their right index finger for 30 s during tACS (70 Hz) over the left M1 and the right cerebellar hemisphere. Each subject participated in 10 trials, separated by intervals of 3 min. Three stimulation conditions were applied: (1) pseudo-stimulation (sham condition), (2) tACS with a current intensity of 1.0 mA and (3) tACS with a current intensity of 2.0 mA. Our results indicated that regardless of the current intensity, participants with the lower motor performance had better motor performance under both tACS conditions. Such a correlation was not observed under the sham condition. Collectively, our findings demonstrated that the effect of gamma tACS over M1 and the cerebellar hemisphere does not depend on current intensity, and that motor performance slightly improved in both tACS conditions for participants with lower motor performance.

Gamma tACS over M1 and cerebellar hemisphere improves motor performance in a phase-specific manner

Transcranial alternating current stimulation (tACS) modulates neural connectivity in the cortical area under the electrodes. Although gamma tACS over the M1 and the cerebellar hemisphere has been shown to improve motor performance, the details of this effect remain unclear. This study aimed to clarify whether the effect of gamma tACS over the M1 and the cerebellar hemisphere is a phase-specific. We applied tACS at 70 Hz over the left M1 and the right cerebellar hemisphere to 20 healthy adults while they performed a visuomotor control task using their right index finger for 30 s. Three stimulation conditions were applied: (1) pseudo-stimulation (sham condition), (2) 180° phase difference in the two cortical target areas (anti-phase condition) and (3) 0° phase difference (in-phase condition). The anti-phase condition decreased task error compared to the sham condition (P = 0.021), but did not differ from the in-phase condition. Our study demonstrated that the effect of gamma tACS over the M1 and the cerebellar hemisphere has phase specificity. This result suggests that intercortical functional synchronization is involved in the improvement of motor performance during gamma tACS over the M1 and the cerebellar hemisphere.

Enhanced action performance following TMS manipulation of associative plasticity in ventral premotor-motor pathway

Skillful goal-directed manual actions such as grasping and manipulating objects are supported by a large sensorimotor network. Within this network, the ventral premotor cortex (PMv) transforms visual information about objects into motor commands that are conveyed to the primary motor cortex (M1), allowing fine control of finger movements. However, it is unknown whether transcranial magnetic stimulation (TMS) of this PMv-to-M1 hierarchical pathway improves action performance. To fill in this gap, here, we used cortico-cortical paired associative stimulation (ccPAS) with the aim of manipulating synaptic efficacy in the human PMv-to-M1 pathway. We found that repeatedly pairing TMS of pre-and post-synaptic nodes of the PMv-to-M1 pathway (i.e., PMv-to-M1 ccPAS) increased motor excitability and enhanced performance on the 9-Hole Peg Test (9-HPT), which taps into PMv-M1 functioning. These effects were specific to the ccPAS protocol consistent with the direction of the PMv-to-M1 hierarchy, as no effects were observed when reversing the order of the paired TMS pulses (i.e., following a M1-to-PMv ccPAS) or when administering sham ccPAS. Additionally, the effect of PMv-to-M1 ccPAS appeared functionally specific, as no behavioral enhancement was observed in a visuomotor control task. We therefore provide novel causal evidence that the PMv-to-M1 pathway, which is instrumental to object-oriented hand actions, is sensitive to TMS manipulations of associative plasticity. Our study highlights the causal role of the PMv-to-M1 pathway in controlling skillful object-oriented hand actions and suggests that ccPAS might be a useful tool for investigating the functional relevance of directional connectivity in humans. These findings may have implications for designing novel therapeutic strategies based on the manipulation of associative plasticity in cortico-cortical networks.

Orofacial and thumb-index finger ramp-and-hold isometric force dynamics in young neurotypical adults

The relation among several parameters of the ramp-and-hold isometric force contraction (peak force and dF/dtmax during the initial phase of force recruitment, and the proportion of hold-phase at target) was quantified for the right and left thumb-index finger pinch, and lower lip midline compression in 40 neurotypical right-handed young adults (20 female/20 males) using wireless force sensors and data acquisition technology developed in our laboratory. In this visuomotor control task, participants produced ramp-and-hold isometric forces as ‘rapidly and accurately’ as possible to end-point target levels at 0.25, 0.5, 1 and 2 Newtons presented to a computer monitor in a randomized block design. Significant relations were found between the parameters of the ramp-and-hold lip force task and target force level, including the peak rate of force change (dF/dtmax), peak force, and the criterion percentage of force within ±5% of target during the contraction hold phase. A significant performance advantage was found among these force variables for the thumb-index finger over the lower lip. The maximum voluntary compression force (MVCF) task revealed highly significant differences in force output between the thumb-index fingers and lower lip (∼4.47–4.70 times greater for the digits versus lower lip), a significant advantage of the right thumb-index finger over the non-dominant left thumb-index finger (12% and 25% right hand advantage for males and females, respectively), and a significant sex difference (∼1.65–1.73 times greater among males).

Diverse coordinate frames on sensorimotor areas in visuomotor transformation

The visuomotor transformation during a goal-directed movement may involve a coordinate transformation from visual ‘extrinsic’ to muscle-like ‘intrinsic’ coordinate frames, which might be processed via a multilayer network architecture composed of neural basis functions. This theory suggests that the postural change during a goal-directed movement task alters activity patterns of the neurons in the intermediate layer of the visuomotor transformation that recieves both visual and proprioceptive inputs, and thus influence the multi-voxel pattern of the blood oxygenation level dependent signal. Using a recently developed multi-voxel pattern decoding method, we found extrinsic, intrinsic and intermediate coordinate frames along the visuomotor cortical pathways during a visuomotor control task. The presented results support the hypothesis that, in human, the extrinsic coordinate frame was transformed to the muscle-like frame over the dorsal pathway from the posterior parietal cortex and the dorsal premotor cortex to the primary motor cortex.

Pseudoneglect in line bisection judgement is associated with a modulation of right hemispheric spatial attention dominance in right-handers

The objective of this study was to validate a line bisection judgement (LBJ) task for use in investigating the lateralized cerebral bases of spatial attention in a sample of 51 right-handed healthy participants. Using functional magnetic resonance imaging (fMRI), the participants performed a LBJ task that was compared to a visuomotor control task during which the participants made similar saccadic and motoric responses. Cerebral lateralization was determined using a voxel-based functional asymmetry analysis and a hemispheric functional lateralization index (HFLI) computed from fMRI contrast images. Behavioural attentional deviation biases were assessed during the LBJ task and a “paper and pencil” symbol cancellation task (SCT). Individual visuospatial skills were also evaluated. The results showed that both the LBJ and SCT tasks elicited leftward spatial biases in healthy subjects, although the biases were not correlated, which indicated their independence. Neuroimaging results showed that the LBJ task elicited a right hemispheric lateralization, with rightward asymmetries found in a large posterior occipito-parietal area, the posterior calcarine sulcus (V1p) and the temporo-occipital junction (TOJ) and in the inferior frontal gyrus, the anterior insula and the superior medial frontal gyrus. The comparison of the LBJ asymmetry map to the lesion map of neglect patients who suffer line bisection deviation demonstrated maximum overlap in a network that included the middle occipital gyrus (MOG), the TOJ, the anterior insula and the inferior frontal region, likely subtending spatial LBJ bias. Finally, the LBJ task-related cerebral lateralization was specifically correlated with the LBJ spatial bias but not with the SCT bias or with the visuospatial skills of the participants. Taken together, these results demonstrated that the LBJ task is adequate for investigating spatial lateralization in healthy subjects and is suitable for determining the factors underlying the variability of spatial cerebral lateralization.

Sleep spindle-related reactivation of category-specific cortical regions after learning face-scene associations

Newly acquired declarative memory traces are believed to be reactivated during NonREM sleep to promote their hippocampo-neocortical transfer for long-term storage. Yet it remains a major challenge to unravel the underlying neuronal mechanisms. Using simultaneous electroencephalography (EEG) and functional magnetic resonance imaging (fMRI) recordings in humans, we show that sleep spindles play a key role in the reactivation of memory-related neocortical representations. On separate days, participants either learned face-scene associations or performed a visuomotor control task. Spindle-coupled reactivation of brain regions representing the specific task stimuli was traced during subsequent NonREM sleep with EEG-informed fMRI. Relative to the control task, learning face-scene associations triggered a stronger combined activation of neocortical and hippocampal regions during subsequent sleep. Notably, reactivation did not only occur in temporal synchrony with spindle events but was tuned by ongoing variations in spindle amplitude. These learning-related increases in spindle-coupled neocortical activity were topographically specific because reactivation was restricted to the face- and scene-selective visual cortical areas previously activated during pre-sleep learning. Spindle-coupled hippocampal activation was stronger the better the participant had performed at prior learning. These results are in agreement with the notion that sleep spindles orchestrate the reactivation of new hippocampal-neocortical memories during sleep.

Quiet Eye Training in a Visuomotor Control Task

Several researchers have reported the importance of maintaining a longer final fixation on the target (termed the quiet eye period, QE) before performing an aiming task. We present an innovative, perceptual training intervention intended to improve the efficiency of gaze behavior (i.e., QE) in shotgun shooting. A sample of 20 international-level skeet shooters were assigned equally to one of two ability-matched groups based on their pretest shooting scores. A perceptual training group participated in a four-step preshot routine alongside three video feedback sessions involving their own gaze behaviors and those of an expert model in an effort to positively influence QE behaviors. A control group received video feedback of performance but without the addition of feedback on QE behaviors. Participants completed pretests and posttests along with an 8-wk training intervention. Subjects of the perceptual training group significantly increased their mean QE duration (397 vs 423 ms), used an earlier onset of QE (257 vs 244 ms), and recorded higher shooting accuracy scores (62 vs 70%) from pretest to posttest. Participants in the perceptual training group significantly reduced gun barrel displacement and absolute peak velocity on the posttest compared with the pretest, although neither variable was overtly trained. A transfer test based on performance during competition indicated that perceptual training significantly improved shooting accuracy from before to after the intervention. No pretest to posttest differences were observed for the control group on the measures reported. The results demonstrate the effectiveness of QE training in improving shooting accuracy and developing a more efficient visuomotor control strategy.The findings have implications for future research on training visuomotor behaviors, attention, and gaze orientation during the performance of aiming tasks.

Cognitive processing and functional interactions within hippocampal-neocortical networks during a spatial working memory task in humans

Cognitive processing and functional interactions within hippocampal-neocortical networks during a spatial working memory task in humans

Cognitive deficits and striato-frontal dopamine release in Parkinson's disease

Idiopathic Parkinson's disease (PD) is often accompanied by a pattern of executive deficits similar to those found in patients with frontal lobe lesions. We investigated whether such cognitive deficits are attributable to frontal lobe dysfunction as a direct consequence of impaired mesocortical dopaminergic transmission or an indirect consequence of impaired nigrostriatal dopaminergic function. For this purpose, changes in synaptic dopamine levels during task performance were monitored using a marker of dopamine D2-receptor availability (11)C-raclopride (RAC) PET. During RAC PET, seven patients with early symptomatic PD and seven age-matched healthy controls performed two types of behavioural task, a spatial working memory task (SWT) and a visuomotor control task (VMT). The SWT involves an executive process which is known to be impaired by both frontal lobe lesions and PD while the VMT is a control test for the visuomotor component of the SWT. Parametric images of RAC binding potential during performance of each task were generated, and compared between the tasks using voxel-based statistical parametric mapping as well as region of interest analysis. In controls, RAC binding was reduced in the dorsal caudate during performance of the SWT compared with the VMT, compatible with increased levels of endogenous dopamine release due to the executive process. In PD patients, this RAC binding reduction was not observed. In contrast, RAC binding in the anterior cingulate cortex within the medial prefrontal cortex was reduced by a comparable level during the SWT both in controls and PD patients. Statistical comparisons between controls and PD patients confirmed significantly attenuated dopamine release in the dorsal caudate in PD, but preserved levels of medial prefrontal dopamine release. Our data suggest that executive deficits in early patients with PD are associated with impaired nigrostriatal dopaminergic function resulting in abnormal processing in the cortico-basal ganglia circuit. In contrast, mesocortical dopaminergic transmission appears well preserved in early PD patients.

Increase in reaction time for solving problems during learning-set formation

Six rhesus monkeys were tested for a change in reaction time for problem-solving during a learning-set task, in which they showed progressive improvement in the rate of learning successive problems of visual discrimination. To evaluate the processing time for cognitive processes in problem-solving, the differences in release latency and movement time between the visual discrimination task and the visuomotor control task were defined. In their first experience, the monkeys required several hundreds of trials for solving the problem, and the Δrelease latency was constant throughout the learning. With increasing experience, they solved problems within fewer trials than with the first problem. At this stage, the Δrelease latency was high at the beginning and then decreased. The rise in the Δrelease latency within the learning acquisition period increased depending on the amount of experience with problems they had solved, whereas the Δmovement time within that period was not significantly affected by the experience with problems. The present findings suggest that the number of problem-solving experiences could promote profound cognitive processing, which may be related to a conceptual representation that actualizes the flexibility of learning, namely, the learning set.

Dopaminergic modulation of high-level cognition in Parkinson's disease: the role of the prefrontal cortex revealed by PET.

This study examined the effects of L-dopa medication in patients with Parkinson's disease on cortical and subcortical blood flow changes during two tasks known to involve frontostriatal circuitry. Eleven patients with Parkinson's disease were scanned on two occasions, one ON L-dopa medication and one OFF L-dopa medication, during performance of the Tower of London planning task and a related test that emphasized aspects of spatial working memory. L-dopa-induced decreases were observed in the right dorsolateral prefrontal cortex during performance of both the planning and the spatial working memory tasks compared with the visuomotor control task. Conversely, L-dopa-induced blood flow increases were observed in the right occipital lobe during the memory task relative to the control task. Data from age-matched healthy volunteers demonstrated that L-dopa effectively normalized blood flow in these regions in the patient group. Moreover, a significant correlation was found between L-dopa-induced, planning related blood flow decreases in the right dorsolateral prefrontal cortex and L-dopa-induced changes in performance on the planning task. These data suggest that L-dopa ameliorates high-level cognitive deficits in Parkinson's disease by inducing relative blood flow changes in the right dorsolateral prefrontal cortex.

Imaging the mental components of a planning task

The Tower of London task (TOL) has been widely used to assess the ability to plan. We used H 2O 15-positron emission tomography to isolate some of the cognitive components of the task. Ten male volunteers were scanned twice in each of six conditions. In two conditions (plan) the subjects had to plan the best solution to TOL problems. In two other conditions (plan–control) the subjects were required to generate four moves without being constrained by a goal. In plan and plan–control tasks the subjects either planned the moves and then executed them (MOVE conditions) or imagined the necessary moves (IMAGINE conditions). The plan and plan–control tasks were matched for the working memory load and ‘initial thinking time’. A visuomotor control task and rest served as baseline conditions. Performance on the plan tasks, in contrast to the baseline conditions, was associated with activation in the dorsal prefrontal cortex, premotor and parietal cortex, and cerebellum. Performance of the plan–control tasks was associated with activation of the same areas. Contrasting the plan with the plan–control tasks revealed no residual activation in the prefrontal cortex. These data show that the activity of the dorsolateral prefrontal cortex on the TOL can be accounted for by the components of generating, selecting and/or remembering mental moves. The task of relating the moves to the goal involves a comparison with a representation of the goal in posterior association areas. We did not find evidence that activation of the dorsal prefrontal cortex is specifically related to the evaluation of a path towards a specified goal, a key component of planning.