Selective attention and decision-making have separable neural bases in space and time.

Attention and decision-making processes are fundamental to cognition. However, they are usually experimentally confounded, making it difficult to link neural observations to specific processes. Here we separated the effects of selective attention from the effects of decision-making on brain activity obtained from human participants (both sexes), using a two-stage task where the attended stimulus and decision were orthogonal and separated in time. Multivariate pattern analyses of multimodal neuroimaging data revealed the dynamics of perceptual and decision-related information coding through time (magnetoencephalography (MEG)), space (functional Magnetic Resonance Imaging (fMRI)), and their combination (MEG-fMRI fusion). Our MEG results showed an effect of attention before decision-making could begin, and fMRI results showed an attention effect in early visual and frontoparietal regions. Model-based MEG-fMRI fusion suggested that attention boosted stimulus information in frontoparietal and early visual regions before decision-making was possible. Together, our results suggest that attention affects neural stimulus representations in frontoparietal regions independent of decision-making.Significance statement Attention and decision-making processes are often experimentally confounded in neuroimaging studies, as participants are commonly asked to make categorical decisions about an attended stimulus only. Our study addresses this issue by separating the effects of selective attention from decision-making effects in human observers. We used multivariate pattern analyses to investigate the dynamics of perceptual and decision-related information coding through time (with MEG) and space (with fMRI) and applied a MEG-fMRI fusion analysis to combine data across neuroimaging modalities. Our results show that attention boosts stimulus information in frontoparietal and early visual regions before decision-making was possible. These findings provide an important verification of claims that attention modulates information processing in the brain and highlights the importance of separating these processes.

Temporal dissociation of neural activity underlying synesthetic and perceptual colors.

Grapheme-color synesthetes experience color when seeing achromatic symbols. We examined whether similar neural mechanisms underlie color perception and synesthetic colors using magnetoencephalography. Classification models trained on neural activity from viewing colored stimuli could distinguish synesthetic color evoked by achromatic symbols after a delay of ∼100 ms. Our results provide an objective neural signature for synesthetic experience and temporal evidence consistent with higher-level processing in synesthesia.

Vicarious touch: Overlapping neural patterns between seeing and feeling touch.

Simulation theories propose that vicarious touch arises when seeing someone else being touched triggers corresponding representations of being touched. Prior electroencephalography (EEG) findings show that seeing touch modulates both early and late somatosensory responses (measured with or without direct tactile stimulation). Functional Magnetic Resonance Imaging (fMRI) studies have shown that seeing touch increases somatosensory cortical activation. These findings have been taken to suggest that when we see someone being touched, we simulate that touch in our sensory systems. The somatosensory overlap when seeing and feeling touch differs between individuals, potentially underpinning variation in vicarious touch experiences. Increases in amplitude (EEG) or cerebral blood flow response (fMRI), however, are limited in that they cannot test for the information contained in the neural signal: seeing touch may not activate the same information as feeling touch. Here, we use time-resolved multivariate pattern analysis on whole-brain EEG data from people with and without vicarious touch experiences to test whether seen touch evokes overlapping neural representations with the first-hand experience of touch. Participants felt touch to the fingers (tactile trials) or watched carefully matched videos of touch to another person's fingers (visual trials). In both groups, EEG was sufficiently sensitive to allow decoding of touch location (little finger vs. thumb) on tactile trials. However, only in individuals who reported feeling touch when watching videos of touch could a classifier trained on tactile trials distinguish touch location on visual trials. This demonstrates that, for people who experience vicarious touch, there is overlap in the information about touch location held in the neural patterns when seeing and feeling touch. The timecourse of this overlap implies that seeing touch evokes similar representations to later stages of tactile processing. Therefore, while simulation may underlie vicarious tactile sensations, our findings suggest this involves an abstracted representation of directly felt touch.

Linking the Brain with Behavior: The Neural Dynamics of Success and Failure in Goal-directed Behavior.

The human brain is extremely flexible and capable of rapidly selecting relevant information in accordance with task goals. Regions of frontoparietal cortex flexibly represent relevant task information such as task rules and stimulus features when participants perform tasks successfully, but less is known about how information processing breaks down when participants make mistakes. This is important for understanding whether and when information coding recorded with neuroimaging is directly meaningful for behavior. Here, we used magnetoencephalography to assess the temporal dynamics of information processing and linked neural responses with goal-directed behavior by analyzing how they changed on behavioral error. Participants performed a difficult stimulus-response task using two stimulus-response mapping rules. We used time-resolved multivariate pattern analysis to characterize the progression of information coding from perceptual information about the stimulus, cue and rule coding, and finally, motor response. Response-aligned analyses revealed a ramping up of perceptual information before a correct response, suggestive of internal evidence accumulation. Strikingly, when participants made a stimulus-related error, and not when they made other types of errors, patterns of activity initially reflected the stimulus presented, but later reversed, and accumulated toward a representation of the "incorrect" stimulus. This suggests that the patterns recorded at later time points reflect an internally generated stimulus representation that was used to make the (incorrect) decision. These results illustrate the orderly and overlapping temporal dynamics of information coding in perceptual decision-making and show a clear link between neural patterns in the late stages of processing and behavior.

The Influence of Object-Color Knowledge on Emerging Object Representations in the Brain.

The ability to rapidly and accurately recognize complex objects is a crucial function of the human visual system. To recognize an object, we need to bind incoming visual features, such as color and form, together into cohesive neural representations and integrate these with our preexisting knowledge about the world. For some objects, typical color is a central feature for recognition; for example, a banana is typically yellow. Here, we applied multivariate pattern analysis on time-resolved neuroimaging (MEG) data to examine how object-color knowledge affects emerging object representations over time. Our results from 20 participants (11 female) show that the typicality of object-color combinations influences object representations, although not at the initial stages of object and color processing. We find evidence that color decoding peaks later for atypical object-color combinations compared with typical object-color combinations, illustrating the interplay between processing incoming object features and stored object knowledge. Together, these results provide new insights into the integration of incoming visual information with existing conceptual object knowledge.SIGNIFICANCE STATEMENT To recognize objects, we have to be able to bind object features, such as color and shape, into one coherent representation and compare it with stored object knowledge. The MEG data presented here provide novel insights about the integration of incoming visual information with our knowledge about the world. Using color as a model to understand the interaction between seeing and knowing, we show that there is a unique pattern of brain activity for congruently colored objects (e.g., a yellow banana) relative to incongruently colored objects (e.g., a red banana). This effect of object-color knowledge only occurs after single object features are processed, demonstrating that conceptual knowledge is accessed relatively late in the visual processing hierarchy.

Reward Rapidly Enhances Visual Perception.

Rewards exert a deep influence on our cognition and behavior. Here, we used a paradigm in which reward information was provided at either encoding or retrieval of a brief, masked stimulus to show that reward can also rapidly modulate perceptual encoding of visual information. Experiment 1 (n = 30 adults) showed that participants' response accuracy was enhanced when a to-be-encoded grating signaled high reward relative to low reward, but only when the grating was presented very briefly and participants reported that they were not consciously aware of it. Experiment 2 (n = 29 adults) showed that there was no difference in participants' response accuracy when reward information was instead provided at the stage of retrieval, ruling out an explanation of the reward-modulation effect in terms of differences in motivated retrieval. Taken together, our findings provide behavioral evidence consistent with a rapid reward modulation of visual perception, which may not require consciousness.

Seeing versus knowing: The temporal dynamics of real and implied colour processing in the human brain.

Colour is a defining feature of many objects, playing a crucial role in our ability to rapidly recognise things in the world around us and make categorical distinctions. For example, colour is a useful cue when distinguishing lemons from limes or blackberries from raspberries. That means our representation of many objects includes key colour-related information. The question addressed here is whether the neural representation activated by knowing that something is red is the same as that activated when we actually see something red, particularly in regard to timing. We addressed this question using neural timeseries (magnetoencephalography, MEG) data to contrast real colour perception and implied object colour activation. We applied multivariate pattern analysis (MVPA) to analyse the brain activation patterns evoked by colour accessed via real colour perception and implied colour activation. Applying MVPA to MEG data allows us here to focus on the temporal dynamics of these processes. Male and female human participants (N = 18) viewed isoluminant red and green shapes and grey-scale, luminance-matched pictures of fruits and vegetables that are red (e.g., tomato) or green (e.g., kiwifruit) in nature. We show that the brain activation pattern evoked by real colour perception is similar to implied colour activation, but that this pattern is instantiated at a later time. These results suggest that a common colour representation can be triggered by activating object representations from memory and perceiving colours.

Limits on visual awareness of object targets in the context of other object category masks: Investigating bottlenecks in the continuous flash suppression paradigm with hand and tool stimuli.

The continuous flash suppression (CFS) task can be used to investigate what limits our capacity to become aware of visual stimuli. In this task, a stream of rapidly changing mask images to one eye initially suppresses awareness for a static target image presented to the other eye. Several factors may determine the breakthrough time from mask suppression, one of which is the overlap in representation of the target/mask categories in higher visual cortex. This hypothesis is based on certain object categories (e.g., faces) being more effective in blocking awareness of other categories (e.g., buildings) than other combinations (e.g., cars/chairs). Previous work found mask effectiveness to be correlated with category-pair high-level representational similarity. As the cortical representations of hands and tools overlap, these categories are ideal to test this further as well as to examine alternative explanations. For our CFS experiments, we predicted longer breakthrough times for hands/tools compared to other pairs due to the reported cortical overlap. In contrast, across three experiments, participants were generally faster at detecting targets masked by hands or tools compared to other mask categories. Exploring low-level explanations, we found that the category average for edges (e.g., hands have less detail compared to cars) was the best predictor for the data. This low-level bottleneck could not completely account for the specific category patterns and the hand/tool effects, suggesting there are several levels at which object category-specific limits occur. Given these findings, it is important that low-level bottlenecks for visual awareness are considered when testing higher-level hypotheses.

Decoding Digits and Dice with Magnetoencephalography: Evidence for a Shared Representation of Magnitude.

Numerical format describes the way magnitude is conveyed, for example, as a digit ("3") or Roman numeral ("III"). In the field of numerical cognition, there is an ongoing debate of whether magnitude representation is independent of numerical format. Here, we examine the time course of magnitude processing when using different symbolic formats. We presented participants with a series of digits and dice patterns corresponding to the magnitudes of 1 to 6 while performing a 1-back task on magnitude. Magnetoencephalography offers an opportunity to record brain activity with high temporal resolution. Multivariate pattern analysis applied to magnetoencephalographic data allows us to draw conclusions about brain activation patterns underlying information processing over time. The results show that we can cross-decode magnitude when training the classifier on magnitude presented in one symbolic format and testing the classifier on the other symbolic format. This suggests a similar representation of these numerical symbols. In addition, results from a time generalization analysis show that digits were accessed slightly earlier than dice, demonstrating temporal asynchronies in their shared representation of magnitude. Together, our methods allow a distinction between format-specific signals and format-independent representations of magnitude showing evidence that there is a shared representation of magnitude accessed via different symbols.

Multisensory temporal processing in own-body contexts: plausibility of hand ownership does not improve visuo-tactile asynchrony detection.

Tracking one's own body is essential for environmental interaction, and involves integrating multisensory cues with stored information about the body's typical features. Exactly how multisensory information is integrated in own-body perception is still unclear. For example, Ide and Hidaka (Exp Brain Res 228:43-50, 2013) found that participants made less precise visuo-tactile temporal order judgments (TOJ) when viewing hands in a plausible orientation (upright; typical for one's own hand) compared to an implausible orientation (rotated 180°). This suggests that viewing one's own body relaxes the precision for perceived visuo-tactile synchrony. In contrast, visuo-proprioceptive research shows improvements for multisensory temporal perception near one's own body in asynchrony detection tasks, implying an increase in precision. Hence, it is unclear whether viewed hand orientation generally modulates the ability to detect small asynchronies between vision and touch, or if this effect is specific to TOJ tasks. We investigated whether viewed hand orientation affects detection of visuo-tactile asynchrony. In two experiments, participants viewed model hands in anatomically plausible or implausible orientations. In one experiment, we stroked the hands to induce the rubber hand illusion. Participants were asked to detect short delays (40-280 ms) between vision (an LED flash on the model hand) and touch (a tap to fingertip of the participant's hidden hand) in a two-interval forced-choice task. Bayesian analyses show that our data provide strong evidence that viewed hand orientation does not affect visuo-tactile asynchrony detection. This study suggests the mechanisms for fine-grained time perception differ between visuo-tactile and visuo-proprioceptive contexts.

More is better: Relative prevalence of multiple targets affects search accuracy.

In real-world searches such as airport baggage screening and radiological examinations, miss errors can be life threatening. Misses increase for additional targets after detecting an initial target, termed "subsequent search misses" (SSMs), and also when targets are more often absent than present, termed the low-prevalence effect. Real-world search tasks often contain more than one target, but the prevalence of these multitarget occasions varies. For example, a cancerous tumor sometimes coexists with a benign tumor and sometimes exists alone. This study aims to investigate how the relative prevalence of multiple targets affects search accuracy. Naive observers searched for all Ts (zero, one, or two) among Ls. In Experiment 1, SSMs occurred in small but not large set sizes, which may be explained by classic capacity limit effects such as the attentional blink and repetition blindness. Experiment 2 showed an interaction between SSMs and the relative prevalence of dual-target trials: Low prevalence of dual-target trials increased SSMs relative to high prevalence dual-target trials. The prevalence of dual-target trials did not affect accuracy on single-target trials. These results may provide a novel avenue for reducing misses by increasing the prevalence of instances with multiple targets. Future efforts should take into account the relative prevalence of multiple targets to effectively reduce life-threatening miss errors.

Feature-selective Attention in Frontoparietal Cortex: Multivoxel Codes Adjust to Prioritize Task-relevant Information.

Human cognition is characterized by astounding flexibility, enabling us to select appropriate information according to the objectives of our current task. A circuit of frontal and parietal brain regions, often referred to as the frontoparietal attention network or multiple-demand (MD) regions, are believed to play a fundamental role in this flexibility. There is evidence that these regions dynamically adjust their responses to selectively process information that is currently relevant for behavior, as proposed by the "adaptive coding hypothesis" [Duncan, J. An adaptive coding model of neural function in prefrontal cortex. Nature Reviews Neuroscience, 2, 820-829, 2001]. Could this provide a neural mechanism for feature-selective attention, the process by which we preferentially process one feature of a stimulus over another? We used multivariate pattern analysis of fMRI data during a perceptually challenging categorization task to investigate whether the representation of visual object features in the MD regions flexibly adjusts according to task relevance. Participants were trained to categorize visually similar novel objects along two orthogonal stimulus dimensions (length/orientation) and performed short alternating blocks in which only one of these dimensions was relevant. We found that multivoxel patterns of activation in the MD regions encoded the task-relevant distinctions more strongly than the task-irrelevant distinctions: The MD regions discriminated between stimuli of different lengths when length was relevant and between the same objects according to orientation when orientation was relevant. The data suggest a flexible neural system that adjusts its representation of visual objects to preferentially encode stimulus features that are currently relevant for behavior, providing a neural mechanism for feature-selective attention.

Adaptive Motor Imagery: A Multimodal Study of Immobilization-Induced Brain Plasticity.

The consequences of losing the ability to move a limb are traumatic. One approach that examines the impact of pathological limb nonuse on the brain involves temporary immobilization of a healthy limb. Here, we investigated immobilization-induced plasticity in the motor imagery (MI) circuitry during hand immobilization. We assessed these changes with a multimodal paradigm, using functional magnetic resonance imaging (fMRI) to measure neural activation, magnetoencephalography (MEG) to track neuronal oscillatory dynamics, and transcranial magnetic stimulation (TMS) to assess corticospinal excitability. fMRI results show a significant decrease in neural activation for MI of the constrained hand, localized to sensorimotor areas contralateral to the immobilized hand. MEG results show a significant decrease in beta desynchronization and faster resynchronization in sensorimotor areas contralateral to the immobilized hand. TMS results show a significant increase in resting motor threshold in motor cortex contralateral to the constrained hand, suggesting a decrease in corticospinal excitability in the projections to the constrained hand. These results demonstrate a direct and rapid effect of immobilization on MI processes of the constrained hand, suggesting that limb nonuse may not only affect motor execution, as evidenced by previous studies, but also MI. These findings have important implications for the effectiveness of therapeutic approaches that use MI as a rehabilitation tool to ameliorate the negative effects of limb nonuse.

Flexible Coding of Task Rules in Frontoparietal Cortex: An Adaptive System for Flexible Cognitive Control.

How do our brains achieve the cognitive control that is required for flexible behavior? Several models of cognitive control propose a role for frontoparietal cortex in the structure and representation of task sets or rules. For behavior to be flexible, however, the system must also rapidly reorganize as mental focus changes. Here we used multivoxel pattern analysis of fMRI data to demonstrate adaptive reorganization of frontoparietal activity patterns following a change in the complexity of the task rules. When task rules were relatively simple, frontoparietal cortex did not hold detectable information about these rules. In contrast, when the rules were more complex, frontoparietal cortex showed clear and decodable rule discrimination. Our data demonstrate that frontoparietal activity adjusts to task complexity, with better discrimination of rules that are behaviorally more confusable. The change in coding was specific to the rule element of the task and was not mirrored in more specialized cortex (early visual cortex) where coding was independent of difficulty. In line with an adaptive view of frontoparietal function, the data suggest a system that rapidly reconfigures in accordance with the difficulty of a behavioral task. This system may provide a neural basis for the flexible control of human behavior.

Attention enhances multi-voxel representation of novel objects in frontal, parietal and visual cortices.

Selective attention is fundamental for human activity, but the details of its neural implementation remain elusive. One influential theory, the adaptive coding hypothesis (Duncan, 2001, An adaptive coding model of neural function in prefrontal cortex, Nature Reviews Neuroscience 2:820-829), proposes that single neurons in certain frontal and parietal regions dynamically adjust their responses to selectively encode relevant information. This selective representation may in turn support selective processing in more specialized brain regions such as the visual cortices. Here, we use multi-voxel decoding of functional magnetic resonance images to demonstrate selective representation of attended--and not distractor--objects in frontal, parietal, and visual cortices. In addition, we highlight a critical role for task demands in determining which brain regions exhibit selective coding. Strikingly, representation of attended objects in frontoparietal cortex was highest under conditions of high perceptual demand, when stimuli were hard to perceive and coding in early visual cortex was weak. Coding in early visual cortex varied as a function of attention and perceptual demand, while coding in higher visual areas was sensitive to the allocation of attention but robust to changes in perceptual difficulty. Consistent with high-profile reports, peripherally presented objects could also be decoded from activity at the occipital pole, a region which corresponds to the fovea. Our results emphasize the flexibility of frontoparietal and visual systems. They support the hypothesis that attention enhances the multi-voxel representation of information in the brain, and suggest that the engagement of this attentional mechanism depends critically on current task demands.

Long-term plasticity in adult somatosensory cortex: functional reorganization after surgical removal of an arteriovenous malformation.

The temporal scale of neuroplasticity following acute alterations in brain structure due to neurosurgical intervention is still under debate. We conducted a longitudinal study with the objective of investigating the postoperative changes in a patient who underwent cerebrovascular surgery and who subsequently lost proprioception in the fingers of her right hand. The results show increased activation in contralesional somatosensory areas, additional recruitment of premotor and posterior parietal areas, and changes in functional connectivity with left postcentral gyrus. These findings demonstrate long-term modifications of cortical organization and as such have important implications for treatment strategies for patients with brain injury.

Symptoms of PTSD Associated With Painful and Nonpainful Vicarious Reactivity Following Amputation.

Although the experience of vicarious sensations when observing another in pain have been described postamputation, the underlying mechanisms are unknown. We investigated whether vicarious sensations are related to posttraumatic stress disorder (PTSD) symptoms and chronic pain. In Study 1, 236 amputees completed questionnaires about phantom limb phenomena and vicarious sensations to both innocuous and painful sensory experiences of others. There was a 10.2% incidence of vicarious sensations, which was significantly more prevalent in amputees reporting PTSD-like experiences, particularly increased arousal and reexperiencing the event that led to amputation (φ = .16). In Study 2, 63 amputees completed the Empathy for Pain Scale and PTSD Checklist-Civilian Version. Cluster analyses revealed 3 groups: 1 group did not experience vicarious pain or PTSD symptoms, and 2 groups were vicarious pain responders, but only 1 had increased PTSD symptoms. Only the latter group showed increased chronic pain severity compared with the nonresponder group (p = .025) with a moderate effect size (r = .35). The findings from both studies implicated an overlap, but also divergence, between PTSD symptoms and vicarious pain reactivity postamputation. Maladaptive mechanisms implicated in severe chronic pain and physical reactivity posttrauma may increase the incidence of vicarious reactivity to the pain of others.

A conceptual lemon: theta burst stimulation to the left anterior temporal lobe untangles object representation and its canonical color.

Object recognition benefits greatly from our knowledge of typical color (e.g., a lemon is usually yellow). Most research on object color knowledge focuses on whether both knowledge and perception of object color recruit the well-established neural substrates of color vision (the V4 complex). Compared with the intensive investigation of the V4 complex, we know little about where and how neural mechanisms beyond V4 contribute to color knowledge. The anterior temporal lobe (ATL) is thought to act as a "hub" that supports semantic memory by integrating different modality-specific contents into a meaningful entity at a supramodal conceptual level, making it a good candidate zone for mediating the mappings between object attributes. Here, we explore whether the ATL is critical for integrating typical color with other object attributes (object shape and name), akin to its role in combining nonperceptual semantic representations. In separate experimental sessions, we applied TMS to disrupt neural processing in the left ATL and a control site (the occipital pole). Participants performed an object naming task that probes color knowledge and elicits a reliable color congruency effect as well as a control quantity naming task that also elicits a cognitive congruency effect but involves no conceptual integration. Critically, ATL stimulation eliminated the otherwise robust color congruency effect but had no impact on the numerical congruency effect, indicating a selective disruption of object color knowledge. Neither color nor numerical congruency effects were affected by stimulation at the control occipital site, ruling out nonspecific effects of cortical stimulation. Our findings suggest that the ATL is involved in the representation of object concepts that include their canonical colors.

Spatial attention increases high-frequency gamma synchronisation in human medial visual cortex.

Visual information processing involves the integration of stimulus and goal-driven information, requiring neuronal communication. Gamma synchronisation is linked to neuronal communication, and is known to be modulated in visual cortex both by stimulus properties and voluntarily-directed attention. Stimulus-driven modulations of gamma activity are particularly associated with early visual areas such as V1, whereas attentional effects are generally localised to higher visual areas such as V4. The absence of a gamma increase in early visual cortex is at odds with robust attentional enhancements found with other measures of neuronal activity in this area. Here we used magnetoencephalography (MEG) to explore the effect of spatial attention on gamma activity in human early visual cortex using a highly effective gamma-inducing stimulus and strong attentional manipulation. In separate blocks, subjects tracked either a parafoveal grating patch that induced gamma activity in contralateral medial visual cortex, or a small line at fixation, effectively attending away from the gamma-inducing grating. Both items were always present, but rotated unpredictably and independently of each other. The rotating grating induced gamma synchronisation in medial visual cortex at 30-70 Hz, and in lateral visual cortex at 60-90 Hz, regardless of whether it was attended. Directing spatial attention to the grating increased gamma synchronisation in medial visual cortex, but only at 60-90 Hz. These results suggest that the generally found increase in gamma activity by spatial attention can be localised to early visual cortex in humans, and that stimulus and goal-driven modulations may be mediated at different frequencies within the gamma range.

Multimodal functional imaging of motor imagery using a novel paradigm.

Neuroimaging studies have shown that the neural mechanisms of motor imagery (MI) overlap substantially with the mechanisms of motor execution (ME). Surprisingly, however, the role of several regions of the motor circuitry in MI remains controversial, a variability that may be due to differences in neuroimaging techniques, MI training, instruction types, or tasks used to evoke MI. The objectives of this study were twofold: (i) to design a novel task that reliably invokes MI, provides a reliable behavioral measure of MI performance, and is transferable across imaging modalities; and (ii) to measure the common and differential activations for MI and ME with functional magnetic resonance imaging (fMRI) and magnetoencephalography (MEG). We present a task in which it is difficult to give accurate responses without the use of either motor execution or motor imagery. The behavioral results demonstrate that participants performed similarly on the task when they imagined vs. executed movements and this performance did not change over time. The fMRI results show a spatial overlap of MI and ME in a number of motor and premotor areas, sensory cortices, cerebellum, inferior frontal gyrus, and ventrolateral thalamus. MI uniquely engaged bilateral occipital areas, left parahippocampus, and other temporal and frontal areas, whereas ME yielded unique activity in motor and sensory areas, cerebellum, precuneus, and putamen. The MEG results show a robust event-related beta band desynchronization in the proximity of primary motor and premotor cortices during both ME and MI. Together, these results further elucidate the neural circuitry of MI and show that our task robustly and reliably invokes motor imagery, and thus may prove useful for interrogating the functional status of the motor circuitry in patients with motor disorders.