Adaptive control is reversed between hands after left hemisphere stroke and lost following right hemisphere stroke.

Human motor adaptability is of utmost utility after neurologic injury such as unilateral stroke. For successful adaptive control of movements, the nervous system must learn to correctly identify the source of a movement error and predictively compensate for this error. The current understanding is that in bimanual tasks, this process is flexible such that errors are assigned to, and compensated for, by the limb that is more likely to produce those errors. Here, we tested the flexibility of the error assignment process in right-handed chronic stroke survivors using a bimanual reaching task in which the hands jointly controlled a single cursor. We predicted that the nondominant left hand in neurotypical adults and the paretic hand in chronic stroke survivors will be more responsible for cursor errors and will compensate more within a trial and learn more from trial to trial. We found that in neurotypical adults, the nondominant left hand does compensate more than the right hand within a trial but learns less trial-to-trial. After a left hemisphere stroke, the paretic right hand compensates more than the nonparetic left hand within-trial but learns less trial-to-trial. After a right hemisphere stroke, the paretic left hand neither corrects more within-trial nor learns more trial-to-trial. Thus, adaptive control of visually guided bimanual reaching movements is reversed between hands after the left hemisphere stroke and lost following the right hemisphere stroke. These results indicate that responsibility assignment is not fully flexible but depends on a central mechanism that is lateralized to the right hemisphere.

The complementary dominance hypothesis: a model for remediating the 'good' hand in stroke survivors.

The complementary dominance hypothesis is a novel model of motor lateralization substantiated by decades of research examining interlimb differences in the control of upper extremity movements in neurotypical adults and hemisphere-specific motor deficits in stroke survivors. In contrast to earlier ideas that attribute handedness to the specialization of one hemisphere, our model proposes complementary motor control specializations in each hemisphere. The dominant hemisphere mediates optimal control of limb dynamics as required for smooth and efficient movements, whereas the non-dominant hemisphere mediates impedance control, important for countering unexpected mechanical conditions and achieving steady-state limb positions. Importantly, this model proposes that each hemisphere contributes its specialization to both arms (though with greater influence from either arm's contralateral hemisphere) and thus predicts that lesions to one hemisphere should produce hemisphere-specific motor deficits in not only the contralesional arm, but also the ipsilesional arm of stroke survivors - a powerful prediction now supported by a growing body of evidence. Such ipsilesional arm motor deficits vary with contralesional arm impairment, and thus individuals with little to no functional use of the contralesional arm experience both the greatest impairments in the ipsilesional arm, as well as the greatest reliance on it to serve as the main or sole manipulator for activities of daily living. Accordingly, we have proposed and tested a novel intervention that reduces hemisphere-specific ipsilesional arm deficits and thereby improves functional independence in stroke survivors with severe contralesional impairment.

Symmetry and synchrony of bimanual movements are not predicated on interlimb control coupling.

Previous studies suggest that bimanual coordination recruits neural mechanisms that explicitly couple control of the arms, resulting in symmetric kinematics. However, the higher symmetry for actions that require congruous joint motions compared with noncongruous joint motions calls into question the concept of control coupling as a general policy. An alternative view proposes that codependence might emerge from an optimal feedback controller that minimizes control effort and costs in task performance. Support for this view comes from studies comparing conditions in which both hands move a shared or independent virtual objects. Because these studies have mainly focused on congruous bimanual movements, it remains unclear if kinematic symmetry emerges from such control policies. We now examine movements with congruous or noncongruous joint motions (inertially symmetric or asymmetric, respectively) under shared or independent cursors conditions. We reasoned that if a control policy minimizes kinematic differences between limbs, spatiotemporal symmetry should remain relatively unaffected by inertial asymmetries. As shared tasks reportedly elicit greater interlimb codependence, these conditions should elicit higher bilateral covariance regardless of inertial asymmetries. Our results indicate a robust spatiotemporal symmetry only under inertially symmetric conditions, regardless of cursor condition. We simulated bimanual reaching using an optimal feedback controller with and without explicit costs of kinematic asymmetry, finding that only the latter mirrored our empirical data. Our findings support the hypothesis that bimanual control policies do not include kinematic asymmetry as a cost when it is not demanded by task constraints suggesting that kinematic symmetry depends critically on mechanical movement conditions.NEW & NOTEWORTHY Previously, the control coupling hypothesis and task-dependent control hypothesis have been shown to be robust in the bimanually symmetrical movement, but whether the same policy remains robust in the bimanually asymmetrical movement remains unclear. Here, with evidence from empirical and simulation data, we show that a spatiotemporal symmetry between the arms is not predicated on control coupling, but instead it is predicated on the symmetry of mechanical conditions (e.g. limb inertia) between the arms.

Bilateral arm movements are coordinated via task-dependent negotiations between independent and codependent control, but not by a "coupling" control policy.

Prior research has shown that coordination of bilateral arm movements might be attributed to either control policies that minimize performance and control costs regardless of bilateral symmetry or by control coupling, which activates bilaterally homologous muscles as a single unit to achieve symmetric performance. We hypothesize that independent bimanual control (movements of one arm are performed without influence on the other) and codependent bimanual control (two arms are constrained to move together with high spatiotemporal symmetry) are two extremes on a coordination spectrum that can be negotiated to meet infinite variations in task demands. To better understand and distinguish between these views, we designed a task where minimization of either control costs or asymmetry would yield different patterns of coordination. Participants made bilateral reaches with a shared visual cursor to a midline target. We then covertly varied the gain contribution of either hand to the shared cursor's horizontal position. Across two experiments, we show that bilateral coordination retains high task-dependent sensitivity to subtle visual feedback gain asymmetries applied to the shared cursor. Specifically, we found a change from strong spatial covariation between hands during equal gains to more independent control during asymmetric gains, which occurred rapidly and with high specificity to the dimension of gain manipulation. Furthermore, the extent of spatial covariation was graded to the magnitude of perpendicular gain asymmetry between hands. These findings suggest coordination of bilateral arm movements flexibly maneuvers along a continuous coordination spectrum in a task-dependent manner that cannot be explained by bilateral control coupling.NEW & NOTEWORTHY Minimization of performance and control costs and efferent coupling between bilaterally homologous muscle groups have been separately hypothesized to describe patterns of bimanual coordination. Here, we address whether the mechanisms mediating independent and codependent control between limbs can be weighted for successful task performance. Using bilaterally asymmetric visuomotor gain perturbations, we show bimanual coordination can be characterized as a negotiation along a spectrum between extremes of independent and codependent control, but not efferent control coupling.

Ipsilesional arm training in severe stroke to improve functional independence (IPSI): phase II protocol.

BACKGROUND: We previously characterized hemisphere-specific motor control deficits in the ipsilesional, less-impaired arm of unilaterally lesioned stroke survivors. Our preliminary data indicate these deficits are substantial and functionally limiting in patients with severe paresis. METHODS: We have designed an intervention ("IPSI") to remediate the hemisphere-specific deficits in the ipsilesional arm, using a virtual-reality platform, followed by manipulation training with a variety of real objects, designed to facilitate generalization and transfer to functional behaviors encountered in the natural environment. This is a 2-site (primary site - Penn State College of Medicine, secondary site - University of Southern California), two-group randomized intervention with an experimental group, which receives unilateral training of the ipsilesional arm throughout 3 one-hour sessions per week for 5 weeks, through our Virtual Reality and Manipulation Training (VRMT) protocol. Our control group receives a conventional intervention on the contralesional arm, 3 one-hour sessions per week for 5 weeks, guided by recently released practice guidelines for upper limb rehabilitation in adult stroke. The study aims to include a total of 120 stroke survivors (60 per group) whose stroke was in the territory of the middle cerebral artery (MCA) resulting in severe upper-extremity motor impairments. Outcome measures (Primary: Jebsen-Taylor Hand Function Test, Fugl-Meyer Assessment, Abilhand, Barthel Index) are assessed at five evaluation points: Baseline 1, Baseline 2, immediate post-intervention (primary endpoint), and 3-weeks (short-term retention) and 6-months post-intervention (long-term retention). We hypothesize that both groups will improve performance of the targeted arm, but that the ipsilesional arm remediation group will show greater improvements in functional independence. DISCUSSION: The results of this study are expected to inform upper limb evaluation and treatment to consider ipsilesional arm function, as part of a comprehensive physical rehabilitation strategy that includes evaluation and remediation of both arms. TRIAL REGISTRATION: This study is registered with ClinicalTrials.gov (Registration ID: NCT03634397 ; date of registration: 08/16/2018).

Competition for limited neural resources in older adults leads to greater asymmetry of bilateral movements than in young adults.

We previously demonstrated that lateralization in the neural control of predictive and impedance mechanisms is reflected by interlimb differences in control of bilateral tasks. Aging has been shown to reduce lateralization during unilateral performance, presumably due to greater recruitment of the ipsilateral hemisphere. We now hypothesize that aging-related reduction in the efficiency of neural resources should produce greater behavioral asymmetry during bilateral actions that require hemispheric specialization for each arm. This is because simultaneous control of dominant and nondominant arm function should induce competition for hemisphere-specific resources. To test this hypothesis, we now examine the effect of aging (young, n = 20; old, n = 20) on performance of a mechanically coupled task, in which one arm reaches toward targets while the other arm stabilizes against a spring that connects the two arms. Results indicate better dominant arm reaching performance and better nondominant arm stabilizing performance for both groups. Most notably, limb and joint compliance was lower in the dominant arm, leading to dominant arm deficits in stabilizing performance. Group analysis indicated that older adults showed substantially greater asymmetry in stabilizing against the spring load than did the younger adults. We propose that competition for limited neural resources in older adults is associated with reduced contributions of right hemisphere mechanisms to right-dominant arm stabilizing performance, and thus to greater asymmetry of performance.NEW & NOTEWORTHY We provide evidence for greater asymmetry of interlimb differences in bilateral coordination for stabilizing and preserved asymmetry of reaching with aging. These results provide the first evidence for increased lateralization with aging within the context of a complementary bilateral task.

Left hemisphere damage produces deficits in predictive control of bilateral coordination.

Previous research has demonstrated hemisphere-specific motor deficits in ipsilesional and contralesional unimanual movements in patients with hemiparetic stroke due to MCA infarct. Due to the importance of bilateral motor actions on activities of daily living, we now examine how bilateral coordination may be differentially affected by right or left hemisphere stroke. To avoid the caveat of simply adding unimanual deficits in assessing bimanual coordination, we designed a unique task that requires spatiotemporal coordination features that do not exist in unimanual movements. Participants with unilateral left (LHD) or right hemisphere damage (RHD) and age-matched controls moved a virtual rectangle (bar) from a midline start position to a midline target. Movement along the long axis of the bar was redundant to the task, such that the bar remained in the center of and parallel to an imaginary line connecting each hand. Thus, to maintain midline position of the bar, movements of one hand closer to or further away from the bar midline required simultaneous, but oppositely directed displacements with the other hand. Our findings indicate that left (LHD), but not right (RHD) hemisphere-damaged patients showed poor interlimb coordination, reflected by significantly lower correlations between displacements of each hand along the bar axis. These left hemisphere-specific deficits were only apparent prior to peak velocity, likely reflecting predictive control of interlimb coordination. In contrast, the RHD group bilateral coordination was not significantly different than that of the control group. We conclude that predictive mechanisms that govern bilateral coordination are dependent on left hemisphere mechanisms. These findings indicate that assessment and training in cooperative bimanual tasks should be considered as part of an intervention framework for post-stroke physical rehabilitation.

Cognitive-perceptual load modulates hand selection in left-handers to a greater extent than in right-handers.

Previous studies have proposed that selecting which hand to use for a reaching task appears to be modulated by a factor described as "task difficulty," defined by either the requirement for spatial precision or movement sequences. However, we previously reported that analysis of the movement costs associated with even simple movements plays a major role in hand selection. We further demonstrated, in right-handers, that cognitive-perceptual loading modulates hand selection by interfering with the analysis of such costs. It has been reported that left-handers tend to show less dominant hand bias in selecting which hand to use during reaching. We, therefore, hypothesized that hand selection would be less affected by cognitive-perceptual loading in left-handers than in right-handers. We employed a visual search task that presented different levels of difficulty (cognitive-perceptual load), as established in previous studies. Our findings indicate that left-handed participants tend to show greater modulation of hand selection by cognitive-perceptual loading than right-handers. Left-handers showed lower dominant hand reaction times than right-handers, and greater high-cost movements that reached to extremes of the contralateral workspace under the most difficult task conditions. We previously showed in this task that midline crossing has high-energy and time costs and that they occur more frequently under cognitively demanding conditions. The current study revealed that midline crossing was associated with the lowest reaction times, in both handedness groups. The fact that left-handers showed lower dominant hand reaction times, and a greater number of high-cost cross-midline reaches under the most cognitively demanding conditions suggests that these actions were erroneous.

Handedness results from complementary hemispheric dominance, not global hemispheric dominance: evidence from mechanically coupled bilateral movements.

Two contrasting views of handedness can be described as 1) complementary dominance, in which each hemisphere is specialized for different aspects of motor control, and 2) global dominance, in which the hemisphere contralateral to the dominant arm is specialized for all aspects of motor control. The present study sought to determine which motor lateralization hypothesis best predicts motor performance during common bilateral task of stabilizing an object (e.g., bread) with one hand while applying forces to the object (e.g., slicing) using the other hand. We designed an experimental equivalent of this task, performed in a virtual environment with the unseen arms supported by frictionless air-sleds. The hands were connected by a spring, and the task was to maintain the position of one hand while moving the other hand to a target. Thus the reaching hand was required to take account of the spring load to make smooth and accurate trajectories, while the stabilizer hand was required to impede the spring load to keep a constant position. Right-handed subjects performed two task sessions (right-hand reach and left-hand stabilize; left-hand reach and right-hand stabilize) with the order of the sessions counterbalanced between groups. Our results indicate a hand by task-component interaction such that the right hand showed straighter reaching performance whereas the left hand showed more stable holding performance. These findings provide support for the complementary dominance hypothesis and suggest that the specializations of each cerebral hemisphere for impedance and dynamic control mechanisms are expressed during bilateral interactive tasks. NEW & NOTEWORTHY We provide evidence for interlimb differences in bilateral coordination of reaching and stabilizing functions, demonstrating an advantage for the dominant and nondominant arms for distinct features of control. These results provide the first evidence for complementary specializations of each limb-hemisphere system for different aspects of control within the context of a complementary bilateral task.

Motor Adaptation Deficits in Ideomotor Apraxia.

OBJECTIVES: The cardinal motor deficits seen in ideomotor limb apraxia are thought to arise from damage to internal representations for actions developed through learning and experience. However, whether apraxic patients learn to develop new representations with training is not well understood. We studied the capacity of apraxic patients for motor adaptation, a process associated with the development of a new internal representation of the relationship between movements and their sensory effects. METHODS: Thirteen healthy adults and 23 patients with left hemisphere stroke (12 apraxic, 11 nonapraxic) adapted to a 30-degree visuomotor rotation. RESULTS: While healthy and nonapraxic participants successfully adapted, apraxics did not. Rather, they showed a rapid decrease in error early but no further improvement thereafter, suggesting a deficit in the slow, but not the fast component of a dual-process model of adaptation. The magnitude of this late learning deficit was predicted by the degree of apraxia, and was correlated with the volume of damage in parietal cortex. Apraxics also demonstrated an initial after-effect similar to the other groups likely reflecting the early learning, but this after-effect was not sustained and performance returned to baseline levels more rapidly, consistent with a disrupted slow learning process. CONCLUSIONS: These findings suggest that the early phase of learning may be intact in apraxia, but this leads to the development of a fragile representation that is rapidly forgotten. The association between this deficit and left parietal damage points to a key role for this region in learning to form stable internal representations. (JINS, 2017, 23, 139-149).

Motor Lateralization Provides a Foundation for Predicting and Treating Non-paretic Arm Motor Deficits in Stroke.

Brain lateralization is a ubiquitous feature of neural organization across the vertebrate spectrum. We have developed a model of motor lateralization that attributes different motor control processes to each cerebral hemisphere. This bilateral hemispheric model of motor control has successfully predicted hemisphere-specific motor control and motor learning deficits in the ipsilesional, or non-paretic, arm of patients with unilateral stroke. We now show across large number and range of stroke patients that these motor performance deficits in the non-paretic arm of stroke patients vary with both the side of the lesion, as well as with the severity of contralesional impairment. This last point can be functionally devastating for patients with severe contralesional paresis because for these individuals, performance of upper extremity activities of daily living depends primarily and often exclusively on ipsilesional arm function. We present a pilot study focused on improving the speed and coordination of ipsilesional arm function in a convenience sample of three stroke patients with severe contralesional impairment. Over a three-week period, patients received a total of nine 1.5 h sessions of training that included intense practice of virtual reality and real-life tasks. Our results indicated substantial improvements in ipsilesional arm movement kinematics, functional performance, and that these improvements carried over to improve functional independence. In addition, the contralesional arm improved in our measure of contralesional impairment, which was likely due to improved participation in activities of daily living. We discuss of our findings for physical rehabilitation.

Preferred directions of arm movements are independent of visual perception of spatial directions.

Directional preferences have previously been demonstrated during horizontal arm movements. These preferences were characterized by a tendency to exploit interaction torques for movement production at the shoulder or elbow, indicating that the preferred directions depend on biomechanical, and not on visual perception-based factors. We directly tested this hypothesis by systematically dissociating visual information from arm biomechanics. Sixteen subjects performed a free-stroke drawing task that required performance of fast strokes from the circle center toward the perimeter, while selecting stroke directions in a random order. Hand position was represented by a cursor displayed in the movement plane. The free-stroke drawing was performed twice, before and after visuomotor adaptation to a 30° clockwise rotation of the perceived hand path. The adaptation was achieved during practicing pointing movements to eight center-out targets. Directional preferences during performance of the free-stroke drawing task were revealed in ten out of the sixteen subjects. The orientation and strength of these preferences were largely the same in both conditions, showing no significant effect of the visuomotor adaptation. In both conditions, the major preferred directions were characterized by higher contribution of interaction torque to net torque at the shoulder as well as by relatively low inertial resistance and the sum of squared shoulder and elbow muscle torques. These results support the hypothesis that directional preferences are largely determined by biomechanical factors. However, this biomechanical effect can decrease or even disappear in some subjects when movements are performed in special conditions, such as the virtual environment used here.

Contralesional motor deficits after unilateral stroke reflect hemisphere-specific control mechanisms.

We have proposed a model of motor lateralization, in which the left and right hemispheres are specialized for different aspects of motor control: the left hemisphere for predicting and accounting for limb dynamics and the right hemisphere for stabilizing limb position through impedance control mechanisms. Our previous studies, demonstrating different motor deficits in the ipsilesional arm of stroke patients with left or right hemisphere damage, provided a critical test of our model. However, motor deficits after stroke are most prominent on the contralesional side. Post-stroke rehabilitation has also, naturally, focused on improving contralesional arm impairment and function. Understanding whether contralesional motor deficits differ depending on the hemisphere of damage is, therefore, of vital importance for assessing the impact of brain damage on function and also for designing rehabilitation interventions specific to laterality of damage. We, therefore, asked whether motor deficits in the contralesional arm of unilateral stroke patients reflect hemisphere-dependent control mechanisms. Because our model of lateralization predicts that contralesional deficits will differ depending on the hemisphere of damage, this study also served as an essential assessment of our model. Stroke patients with mild to moderate hemiparesis in either the left or right arm because of contralateral stroke and healthy control subjects performed targeted multi-joint reaching movements in different directions. As predicted, our results indicated a double dissociation; although left hemisphere damage was associated with greater errors in trajectory curvature and movement direction, errors in movement extent were greatest after right hemisphere damage. Thus, our results provide the first demonstration of hemisphere specific motor control deficits in the contralesional arm of stroke patients. Our results also suggest that it is critical to consider the differential deficits induced by right or left hemisphere lesions to enhance post-stroke rehabilitation interventions.

Hemispheric differences in the control of limb dynamics: a link between arm performance asymmetries and arm selection patterns.

Human handedness has been described and measured from two perspectives: handedness inventories rate hand preferences, whereas other tests examine motor performance asymmetries. These two measurement approaches reflect a major controversy in a literature that defines handedness as either a preference or an asymmetry in sensorimotor processing. Over the past decade, our laboratory has developed a model of handedness based on lateralization of neural processes. This model attributes distinct control processes to each hemisphere, which in turn lead to observable interlimb sensorimotor performance asymmetries. We now hypothesize that arm preference, or choice, may depend on the interaction between sensorimotor performance asymmetries and the given task. The purpose of this study is to examine whether arm selection is linked to interlimb performance asymmetries during reaching. Right-handed subjects made choice and nonchoice reaches to each of eight targets (d = 3.5 cm) arranged radially (r = 13 cm) around a midline starting position. We displaced each cursor (one associated with each hand) 30 cm to the midline start circle to ensure that there were no hemispace-related geometric, mechanical, or perceptual biases to use either arm for the two midline targets. The three targets on each side of the midline received mostly reaches from the ipsilateral arm, a tendency previously described as a "hemispace bias." However, the midline targets, which were equidistant from each hand, received more dominant arm reaches. Dominant arm hand paths to these targets were straighter and more accurately directed. Inverse dynamics analyses revealed a more proficient dominant arm strategy that exploited intersegmental dynamics to a greater extent than did the nondominant arm. These findings suggest that sensorimotor asymmetries in dynamic coordination might explain limb choices. We discuss the implications of these results for theories of action selection, models of handedness, and models of neural lateralization.

Hemispheric specialization for movement control produces dissociable differences in online corrections after stroke.

In this study, we examine whether corrections made during an ongoing movement are differentially affected by left hemisphere damage (LHD) and right hemisphere damage (RHD). Our hypothesis of motor lateralization proposes that control mechanisms specialized to the right hemisphere rely largely on online processes, while the left hemisphere primarily utilizes predictive mechanisms to specify optimal coordination patterns. We therefore predict that RHD, but not LHD, should impair online correction when task goals are unexpectedly changed. Fourteen stroke subjects (7 LHD, 7 RHD) and 14 healthy controls reached to 1 of the 3 targets that unexpectedly "jumped" during movement onset. RHD subjects showed a considerable delay in initiating the corrective response relative to controls and LHD subjects. However, both stroke groups made large final position errors on the target jump trials. Position deficits following LHD were associated with poor intersegmental coordination, while RHD subjects had difficulty terminating their movements appropriately. These findings confirm that RHD, but not LHD, produces a deficit in the timing of online corrections and also indicate that both stroke groups show position deficits that are related to the specialization of their damaged hemisphere. Further research is needed to identify specific neural circuits within each hemisphere critical for these processes.

Left parietal regions are critical for adaptive visuomotor control.

The question addressed in this study is whether parietal brain circuits involved in adaptation to novel visuomotor conditions are lateralized. This information is critical for characterizing the neural mechanisms mediating adaptive behavior in humans, as well as for assessing the effects of unilateral brain damage on function. Moreover, previous research has been controversial in this regard. We compared visuomotor adaptation in 10 patients with focal, unilateral, left or right parietal lesions and healthy control participants. All subjects reached to each of eight targets over three experimental sessions: a baseline session, where the visually displayed and actual hand motion were matched; an adaptation session, where the visual feedback deviated from the actual movement direction by 30°; and an after-effect session, where visual feedback was again matched to hand motion. Adaptation was primarily quantified as a change in initial movement direction throughout the adaptation session and the presence of after-effects when the rotation was removed. Patients with right parietal damage demonstrated normal adaptation and large after-effects, which was comparable to the performance of healthy controls. In contrast, patients with left parietal damage showed a clear deficit in adaptation and showed no after-effects. Thus, our results show that left but not right parietal regions are critical for visuomotor adaptation. These findings are discussed in the context that left parietal regions are critical for the modification of stored representations of the relationship between movement commands and limb and environmental state, as is thought to occur during visuomotor adaptation.

Dynamic dominance varies with handedness: reduced interlimb asymmetries in left-handers.

Our previous studies of interlimb asymmetries during reaching movements have given rise to the dynamic-dominance hypothesis of motor lateralization. This hypothesis proposes that dominant arm control has become optimized for efficient intersegmental coordination, which is often associated with straight and smooth hand-paths, while non-dominant arm control has become optimized for controlling steady-state posture, which has been associated with greater final position accuracy when movements are mechanically perturbed, and often during movements made in the absence of visual feedback. The basis for this model of motor lateralization was derived from studies conducted in right-handed subjects. We now ask whether left-handers show similar proficiencies in coordinating reaching movements. We recruited right- and left-handers (20 per group) to perform reaching movements to three targets, in which intersegmental coordination requirements varied systematically. Our results showed that the dominant arm of both left- and right-handers were well coordinated, as reflected by fairly straight hand-paths and low errors in initial direction. Consistent with our previous studies, the non-dominant arm of right-handers showed substantially greater curvature and large errors in initial direction, most notably to targets that elicited higher intersegmental interactions. While the right, non-dominant, hand-paths of left-handers were slightly more curved than those of the dominant arm, they were also substantially more accurate and better coordinated than the non-dominant arm of right-handers. Our results indicate a similar pattern, but reduced lateralization for intersegmental coordination in left-handers. These findings suggest that left-handers develop more coordinated control of their non-dominant arms than right-handers, possibly due to environmental pressure for right-handed manipulations.

Interlimb differences of directional biases for stroke production.

Directional preferences during center-out horizontal shoulder-elbow movements were previously characterized for the dominant arm. These preferences were attributed to a tendency to actively accelerate one joint, while exploiting largely passive motion at the other joint. Since the non-dominant arm is known for inefficient coordination of inter-segmental dynamics, here we hypothesized that directional preferences would differ between the arms. A center-out free-stroke drawing task was used that allowed freedom in the selection of movement directions. The task was performed both with and without a secondary cognitive task that has been shown to increase directional biases of the dominant arm. Mirror-symmetrical directional preferences were observed in both arms, with similar bias strength and secondary task effects. The preferred directions were characterized by maximal exploitation of interaction torques for movement production, but only in the dominant arm. The non-dominant arm failed to benefit from interaction torques. The results point to a hierarchical architecture of control. At the higher level, a movement capable to perform the task while satisfying preferences in joint control is specified through forward dynamic transformations. This process is mediated for both arms from a common neural network adapted to the dominant arm and, specifically, to its ability to exploit interaction torques. Dynamic transformations that determine actual control commands are specified at the lower level of control. An alternative interpretation that strokes might be planned evenly across directions, and biases emerge during movement execution due to anisotropic resistance of intrinsic factors that do not depend on arm dominance is also discussed.

Critical neural substrates for correcting unexpected trajectory errors and learning from them.

Our proficiency at any skill is critically dependent on the ability to monitor our performance, correct errors and adapt subsequent movements so that errors are avoided in the future. In this study, we aimed to dissociate the neural substrates critical for correcting unexpected trajectory errors and learning to adapt future movements based on those errors. Twenty stroke patients with focal damage to frontal or parietal regions in the left or right brain hemispheres and 20 healthy controls performed a task in which a novel mapping between actual hand motion and its visual feedback was introduced. Only patients with frontal damage in the right hemisphere failed to correct for this discrepancy during the ongoing movement. However, these patients were able to adapt to the distortion such that their movement direction on subsequent trials improved. In contrast, only patients with parietal damage in the left hemisphere showed a clear deficit in movement adaptation, but not in online correction. Left frontal or right parietal damage did not adversely impact upon either process. Our findings thus identify, for the first time, distinct and lateralized neural substrates critical for correcting unexpected errors during ongoing movements and error-based movement adaptation.

Neural Control of Stopping and Stabilizing the Arm.

Stopping is a crucial yet under-studied action for planning and producing meaningful and efficient movements. In this review, we discuss classical human psychophysics studies as well as those using engineered systems that aim to develop models of motor control of the upper limb. We present evidence for a hybrid model of motor control, which has an evolutionary advantage due to division of labor between cerebral hemispheres. Stopping is a fundamental aspect of movement that deserves more attention in research than it currently receives. Such research may provide a basis for understanding arm stabilization deficits that can occur following central nervous system (CNS) damage.