Impairments in action and perception after right intraparietal damage.

We examined visually-guided reaching and perception in an individual who underwent resection of a small tumor in right intraparietal sulcus (pIPS). In the first experiment, she reached to targets presented on a touch screen. Vision was occluded from reach onset on half of the trials, whereas on the other half she had vision during the entire reach. For visually-guided reaching, she demonstrated significantly more reach errors for targets left of fixation versus right of fixation. However, there were no hemispatial differences when reaching without vision. Furthermore, her performance was consistent for reaches with either hand, providing evidence that pIPS encodes location based on an eye-centered reference frame. Second, previous studies reported that optic ataxics are more accurate when reaching to remembered versus visible target locations. We repeated the first experiment, adding a five second delay between stimulus presentation and reach initiation. In contrast to prior reports, she was less accurate in delayed versus immediate reaching. Finally, we examined whether a small pIPS resection would disrupt visuospatial processing in a simple perceptual task. We presented two small circles in succession in either the same location or offset at varying distances, and asked whether the two circles were presented in the same or different position. She was significantly more impaired left of fixation compared to right of fixation, providing evidence for a perceptual deficit after a dorsal stream lesion.

Movement Imitation via an Abstract Trajectory Representation in Dorsal Premotor Cortex.

Humans are particularly good at copying novel and meaningless gestures. The mechanistic and anatomical basis for this specialized imitation ability remains largely unknown. One idea is that imitation occurs by matching body configurations. Here we propose an alternative route to imitation that depends on a body-independent representation of the trajectory path of the end-effector. We studied a group of patients with strokes in the left frontoparietal cortices. We found that they were equally impaired at imitating movement trajectories using the ipsilesional limb (i.e., the nonparetic side) that were cued either by an actor using their whole arm or just by a cursor, suggesting that body configuration information is not always critical for imitation and that a representation of abstract trajectory shape may suffice. In addition, imitation ability was uncorrelated to the ability to identify the trajectory shape, suggesting that imitation deficits were unlikely to arise from perceptual impairments. Finally, a lesion-symptom mapping analysis found that imitation deficits were associated with lesions in left dorsal premotor but not parietal cortex. Together, these findings suggest a novel body-independent route to imitation that relies on the ability to plan abstract movement trajectories within dorsal premotor cortex.SIGNIFICANCE STATEMENT The ability to imitate is critical for rapidly learning to produce new gestures and actions, but how the brain translates observed movements into motor commands is poorly understood. Examining the ability of patients with strokes affecting the left hemisphere revealed that meaningless gestures can be imitated by succinctly representing only the motion of the hand in space, rather than the posture of the entire arm. Moreover, performance deficits correlated with lesions in dorsal premotor cortex, an area not previously associated with impaired imitation of arm postures. These findings thus describe a novel route to imitation that may also be impaired in some patients with apraxia.

The mirror illusion's effects on body state estimation.

The mirror illusion uses a standard mirror to create a compelling illusion in which movements of one limb seem to be made by the other hidden limb. In this paper we adapt a motor control framework to examine which estimates of the body's configuration are affected by the illusion. We propose that the illusion primarily alters estimates related to upcoming states of the body (the desired state and the predicted state), with smaller effects on the estimate of the body state prior to movement initiation. Support for this proposal is provided both by behavioural effects of the illusion and by neuroimaging evidence from one neural region, V6A, that is critically involved in the mirror illusion and limb state estimation more generally.

Corrigendum: The role of action representations in thematic object relations.

[This corrects the article on p. 140 in vol. 8, PMID: 24672461.].

Enhancing the mirror illusion with transcranial direct current stimulation.

Visual feedback has a strong impact on upper-extremity movement production. One compelling example of this phenomena is the mirror illusion (MI), which has been used as a treatment for post-stroke movement deficits (mirror therapy). Previous research indicates that the MI increases primary motor cortex excitability, and this change in excitability is strongly correlated with the mirror's effects on behavioral performance of neurologically-intact controls. Based on evidence that primary motor cortex excitability can also be increased using transcranial direct current stimulation (tDCS), we tested whether bilateral tDCS to the primary motor cortices (anode right-cathode left and anode left-cathode right) would modify the MI. We measured the MI using a previously-developed task in which participants make reaching movements with the unseen arm behind a mirror while viewing the reflection of the other arm. When an offset in the positions of the two limbs relative to the mirror is introduced, reaching errors of the unseen arm are biased by the reflected arm's position. We found that active tDCS in the anode right-cathode left montage increased the magnitude of the MI relative to sham tDCS and anode left-cathode right tDCS. We take these data as a promising indication that tDCS could improve the effect of mirror therapy in patients with hemiparesis.

Effects of vibrotactile feedback on human learning of arm motions.

Tactile cues generated from lightweight, wearable actuators can help users learn new motions by providing immediate feedback on when and how to correct their movements. We present a vibrotactile motion guidance system that measures arm motions and provides vibration feedback when the user deviates from a desired trajectory. A study was conducted to test the effects of vibrotactile guidance on a subject's ability to learn arm motions. Twenty-six subjects learned motions of varying difficulty with both visual (V), and visual and vibrotactile (VVT) feedback over the course of four days of training. After four days of rest, subjects returned to perform the motions from memory with no feedback. We found that augmenting visual feedback with vibrotactile feedback helped subjects reduce the root mean square (rms) angle error of their limb significantly while they were learning the motions, particularly for 1DOF motions. Analysis of the retention data showed no significant difference in rms angle errors between feedback conditions.

The role of action representations in thematic object relations.

A number of studies have explored the role of associative/event-based (thematic) and categorical (taxonomic) relations in the organization of object representations. Recent evidence suggests that thematic information may be particularly important in determining relationships between manipulable artifacts. However, although sensorimotor information is on many accounts an important component of manipulable artifact representations, little is known about the role that action may play during the processing of semantic relationships (particularly thematic relationships) between multiple objects. In this study, we assessed healthy and left hemisphere stroke participants to explore three questions relevant to object relationship processing. First, we assessed whether participants tended to favor thematic relations including action (Th+A, e.g., wine bottle-corkscrew), thematic relationships without action (Th-A, e.g., wine bottle-cheese), or taxonomic relationships (Tax, e.g., wine bottle-water bottle) when choosing between them in an association judgment task with manipulable artifacts. Second, we assessed whether the underlying constructs of event relatedness, action relatedness, and categorical relatedness determined the choices that participants made. Third, we assessed the hypothesis that degraded action knowledge and/or damage to temporo-parietal cortex, a region of the brain associated with the representation of action knowledge, would reduce the influence of action on the choice task. Experiment 1 showed that explicit ratings of event, action, and categorical relatedness were differentially predictive of healthy participants' choices, with action relatedness determining choices between Th+A and Th-A associations above and beyond event and categorical ratings. Experiment 2 focused more specifically on these Th+A vs. Th-A choices and demonstrated that participants with left temporo-parietal lesions, a brain region known to be involved in sensorimotor processing, were less likely than controls and tended to be less likely than patients with lesions sparing that region to use action relatedness in determining their choices. These data indicate that action knowledge plays a critical role in processing of thematic relations for manipulable artifacts.

Response interference between functional and structural object-related actions is increased in patients with ideomotor apraxia.

We report data from two left hemisphere stroke patients with moderate-to-severe ideomotor apraxia who exhibited deficits in positioning their hands to use 'conflict' objects (objects grasped and used with different hand postures) relative to controls and patients with mild apraxia. These novel data support the claim that actions to common objects are subject to interference between multiple responses, and suggest that errors in apraxia may be attributed to deficient resolution of competition between appropriate and inappropriate actions.

Visuo-motor gain adaptation and generalization following left hemisphere stroke.

During gain adaptation, participants must learn to adapt to novel visuo-motor mappings in which the movement amplitudes they produce do not match the visual feedback they receive. The aim of the present study was to investigate the neural substrates of gain adaptation by examining its possible disruption following left hemisphere stroke. Thirteen chronic left hemisphere stroke patients and five healthy right-handed control subjects completed three experimental phases involving reaching with the left hand, which was the less-affected hand in patients. First, participants reached without visual feedback to six different target locations (baseline phase). Next, in the adaptation phase, participants executed movements to one target under conditions in which the perceived movement distance was 70% of the produced movement distance. Last, in order to test the generalization of this new visuomotor mapping, participants made movements without visual feedback to untrained target locations (generalization phase). Significant between-patient differences were observed during adaptation. Lesion analyses indicated that these between-patient differences were predicted by the amount of damage to the supramarginal gyrus (Brodmann area 40). In addition, patients performed more poorly than controls in the generalization phase, suggesting that different processes are involved in adaptation and generalization periods.

Extending Fitts' Law to three-dimensional obstacle-avoidance movements: support for the posture-based motion planning model.

According to Fitts' Law, the time (MT) to move to a target is a linear function of the logarithm of the ratio between the target's distance and width. Although Fitts' Law accurately predicts MTs for direct movements, it does not accurately predict MTs for indirect movements, as when an obstacle intrudes on the direct movement path. To address this limitation, Jax et al. (2007) added an obstacle-intrusion term to Fitts' Law. They accurately predicted MTs around obstacles in two-dimensional (2-D) workspaces, but their model had one more parameter than Fitts' Law did and was merely descriptive. In this study, we addressed these concerns by turning to the mechanistic, posture-based (PB) movement planning model. The PB-based model accounted for almost as much MT variance in a 3-D movement task as the model of Jax et al., with only two parameters, the same number of parameters as in Fitts' Law.

Contributions of efference copy to limb localization: evidence from deafferentation.

Previous research with deafferented subjects suggests that efference copy can be used to update limb position. However, the contributions of efference copy to limb localization are currently unclear. We examined the performance of JDY, a woman with severe, longstanding proprioceptive deficits from a sensory peripheral neuropathy, on a reaching task to explore the contribution of efference copy to trajectory control. JDY and eight healthy controls reached without visual feedback to a target that either remained stationary or jumped to a second location after movement initiation. JDY consistently made hypermetric movements to the final target, exhibiting significant problems with amplitude control. Despite this amplitude control deficit, JDY's performance on jump trials showed that the angle of movement correction (angle between pre- and post-correction movement segments) was significantly correlated with the distance (but not time) of movement from start to turn point. These data suggest that despite an absence of proprioceptive and visual information regarding hand location, JDY derived information about movement distance that informed her movement correction on jump trials. The same type of information that permitted her to correct movement direction on-line, however, was not available for control of final arm position. We propose that efference copy can provide a consistent estimate of limb position that becomes less informative over the course of the movement. We discuss the implications of these data for current models of motor control.

Response interference between functional and structural actions linked to the same familiar object.

Viewing objects with the intention to act upon them may activate task-irrelevant motor responses. Many manufactured objects are associated with two action classes: grasping in accordance with object structure and skillful use consistent with object function. We studied the potential for within-object competition during action selection by comparing initiation latencies for "conflict" objects (with competing structure and function responses) to "non-conflict" objects (with a single response). We demonstrated a novel pattern of within-object interference wherein actions involving conflict objects were slowed when participants skillfully used those objects (grasp-on-use interference) as well as a second pattern of interference when conflict objects were grasped after skillfully using the same objects in previous blocks (long-term use-on-grasp interference). These data suggest that actions to common objects are influenced by competition between rapid but briefly maintained grasp responses and slower but longer-lasting use responses, and advance our understanding of the process and neural substrates of selection for action.

Disorders of the perceptual-motor system.

The study of patients with movement disorders provides insight into both the functional organization and the neural substrates of the perceptual-motor system. By and large, we feel this source of information has been underutilized within the basic science of motor control. To begin to address this shortcoming, this chapter reviews three disorders of the perceptual-motor system (disorders of the body schema, optic ataxia, and ideomotor apraxia) and illustrates how the study of these disorders can inform central issues within the field of motor control. These issues include (1) the need for the perceptual-motor system to maintain a representation of the body's current configuration in order to produce movements, (2) the use of visual information in movement production, (3) the coordinate frame in which movements are controlled, (4) the distinction between movement planning and online correction, and (5) the role of the parietal cortex in action. In the conclusion, we discuss several limitations of studying patients with movement disorders as well as suggest that greater communication is needed between researchers in the basic science of motor control and clinicians developing treatments for movement disorders.

The posture-based motion planning framework: new findings related to object manipulation, moving around obstacles, moving in three spatial dimensions, and haptic tracking.

We describe the results of recent studies inspired by the posture-based motion planning theory (Rosenbaum et al., 2001). The research concerns analyses of human object manipulation, obstacle avoidance, three-dimensional movement generation, and haptic tracking, the findings of which are discussed in relation to whether they support or fail to support the premises of the theory. Each of the aforementioned topics potentially challenges the theory's claim that, in motion, goal postures are planned before the selection of movements towards those postures. However, even the quasi-continuous phenomena under study show features that comply with prospective, end-state-based motion planning. We conclude that progress in motor control should not be frustrated by the view that no model is, or will ever be, optimal. Instead, it should find promise in the steady growth of insights afforded by challenges to existing theories.

Two-component models of reaching: evidence from deafferentation in a Fitts' law task.

Both feedforward and feedback mechanisms are used to ensure accurate movements. Feedback information comes primarily from vision and proprioception; the relative contributions of these modalities to on-line control of action and internal model maintenance remain unclear. We report data from an experiment in which a chronically deafferented subject (JDY) and nine controls were asked to reach to targets of different sizes both with and without vision. Movement times of controls were consistent with Fitts' law on trials with and without vision. JDY's movement times were consistent with Fitts' law only with vision. She was inaccurate relative to controls with vision but exhibited a significantly greater decrement in performance than controls without vision. Finally, JDY's performance on trials with vision deteriorated as a function of the number of preceding trials on which vision was not available. These data provide support for classical models of motor control that divide reaching into an initial ballistic movement guided by efference copy, and a terminal stage where sensory feedback is crucial. Furthermore, these data also demonstrate that proprioception is needed to calibrate and maintain internal models of action.

More than (where the target) meets the eyes: disrupted visuomotor transformations in optic ataxia.

Visually guided reaching entails multiple coordinate frame transformations between retina-centered target location and body-centered limb location. Reaching errors in optic ataxia (OA) may be caused by disruptions to these transformations. Consistent with this proposal, previous studies report that reaching errors in OA depend primarily on the location of a target relative to the patient's gaze regardless of its location relative to the head or body midline. We attempted to replicate this finding by testing KE, a patient with OA following bilateral parietal and left premotor lesions (as well as significant non-specific white matter disease) on a reaching task that varied the orientation of his head and torso while holding the gaze-relative position of the target constant (always foveated). In contrast to previous reports, we observed that rotating the head or body away from the midline led to decreased reaching accuracy. Further analyses showed that multiple visuomotor transformation steps might have been disrupted in KE. These results suggest that gaze-relative target position is not the sole determinant of reaching errors in all OA patients.

Hand path priming in manual obstacle avoidance: rapid decay of dorsal stream information.

The dorsal, action-related, visual stream has been thought to have little or no memory. This hypothesis has seemed credible because functions related to the dorsal stream have been generally unsusceptible to priming from previous experience. Tests of this claim have yielded inconsistent results, however. We argue that these inconsistencies may be due to methodological differences in the time between primes and test stimuli. In this study we sought to clarify the effect of time between primes and test stimuli by having participants complete a visually guided manual obstacle avoidance task with varying times between trials. Consistent with a previous study using this task, we found that hand path curvature depended on the presence or absence of an obstacle in the previous trial. This hand path priming effect decayed quickly as the time between trials increased, and was almost, though not entirely, eliminated when 1000 ms separated successive trials. The results are consistent with the hypothesis that the dorsal stream can be primed but that this priming attenuates rapidly. We suggest that this outcome may indicate that the period over which the dorsal stream retains information may be related to the sequential statistics of action.

Toward an integrated account of object and action selection: a computational analysis and empirical findings from reaching-to-grasp and tool-use.

The act of reaching for and acting upon an object involves two forms of selection: selection of the object as a target, and selection of the action to be performed. While these two forms of selection are logically dissociable, and are evidently subserved by separable neural pathways, they must also be closely coordinated. We examine the nature of this coordination by developing and analyzing a computational model of object and action selection first proposed by Ward [Ward, R. (1999). Interactions between perception and action systems: a model for selective action. In G. W. Humphreys, J. Duncan, & A. Treisman (Eds.), Attention, Space and Action: Studies in Cognitive Neuroscience. Oxford: Oxford University Press]. An interesting tenet of this account, which we explore in detail, is that the interplay between object and action selection depends critically on top-down inputs representing the current task set or plan of action. A concrete manifestation of this, established through a series of simulations, is that the impact of distractor objects on reaching times can vary depending on the nature of the current action plan. In order to test the model's predictions in this regard, we conducted two experiments, one involving direct object manipulation, the other involving tool-use. In both experiments we observed the specific interaction between task set and distractor type predicted by the model. Our findings provide support for the computational model, and more broadly for an interactive account of object and action selection.

Hand path priming in manual obstacle avoidance: evidence for abstract spatiotemporal forms in human motor control.

Previous research suggests that motor equivalence is achieved through reliance on effector-independent spatiotemporal forms. Here the authors report a series of experiments investigating the role of such forms in the production of movement sequences. Participants were asked to complete series of arm movements in time with a metronome and, on some trials, with an obstacle between 1 or more of the target pairs. In moves following an obstacle, participants only gradually reduced the peak heights of their manual jumping movements. This hand path priming effect, scaled with obstacle height, was preserved when participants cleared the obstacle with 1 hand and continued with the other, and it was modulated by future task demands. The results are consistent with the hypothesis that the control of movement sequences relies on abstract spatiotemporal forms. The data also support the view that motor programming is largely achieved by changing just those features that distinguish the next movement to be made from the movement that was just made.

The problem of serial order in behavior: Lashley's legacy.

In a prescient paper Karl Lashley (1951) rejected reflex chaining accounts of the sequencing of behavior and argued instead for a more cognitive account in which behavioral sequences are typically controlled with central plans. An important feature of such plans, according to Lashley, is that they are hierarchical. Lashley offered several sources of evidence for the hierarchical organization for behavioral plans, and others afterward provided more evidence for this hypothesis. We briefly review that evidence here and then shift from a focus on the structure of plans (Lashley's point of concentration) to the processes by which plans are formed in real time. Two principles emerge from the studies we review. One is that plans are not formed from scratch for each successive movement sequence but instead are formed by making whatever changes are needed to distinguish the movement sequence to be performed next from the movement sequence that has just been performed. This plan-modification view is supported by two phenomena discovered in our laboratory: the parameter remapping effect, and the handpath priming effect. The other principle we review is that even single movements appear to be controlled with hierarchically organized plans. At the top level are the starting and goal postures. At the lower level are the intermediate states comprising the transition from the starting posture to the goal posture. The latter principle is supported by another phenomenon discovered in our lab, the end-state comfort effect, and by a computational model of motor planning which accounts for a large number of motor phenomena. Interestingly, the computational model hearkens back to a classical method of generating cartoon animations that relies on the production of keyframes first and the production of interframes (intermediate frames) second.