Differential Effects of Cerebellar Degeneration on Feedforward versus Feedback Control across Speech and Reaching Movements

Errors that result from a mismatch between predicted movement outcomes and sensory afference are used to correct ongoing movements through feedback control and to adapt feedforward control of future movements. The cerebellum has been identified as a critical part of the neural circuit underlying implicit adaptation across a wide variety of movements (reaching, gait, eye movements, and speech). The contribution of this structure to feedback control is less well understood. Although it has recently been shown in the speech domain that individuals with cerebellar degeneration produce larger online corrections for sensory perturbations than control participants, similar behavior has not been observed in other motor domains. Currently, comparisons across domains are limited by different population samples and potential ceiling effects in existing tasks. To assess the relationship between changes in feedforward and feedback control associated with cerebellar degeneration across motor domains, we evaluated adaptive (feedforward) and compensatory (feedback) responses to sensory perturbations in reaching and speech production in human participants of both sexes with cerebellar degeneration and neurobiologically healthy controls. As expected, the cerebellar group demonstrated impaired adaptation in both reaching and speech. In contrast, the groups did not differ in their compensatory response in either domain. Moreover, compensatory and adaptive responses in the cerebellar group were not correlated within or across motor domains. These results point to a general impairment in feedforward control with spared feedback control in cerebellar degeneration. However, the magnitude of feedforward impairments and potential changes in feedback-based control manifest in a domain-specific manner across individuals.SIGNIFICANCE STATEMENT The cerebellum contributes to feedforward updating of movement in response to sensory errors, but its role in feedback control is less understood. Here, we tested individuals with cerebellar degeneration (CD), using sensory perturbations to assess adaptation of feedforward control and feedback gains during reaching and speech production tasks. The results confirmed that CD leads to reduced adaption in both domains. However, feedback gains were unaffected by CD in either domain. Interestingly, measures of feedforward and feedback control were not correlated across individuals within or across motor domains. Together, these results indicate a general impairment in feedforward control with spared feedback control in CD. However, the magnitude of feedforward impairments manifests in a domain-specific manner across individuals.     

Pushing attention to one side: Force field adaptation alters neural correlates of orienting and disengagement of spatial attention

Sensorimotor adaptation to wedge prisms can alter the balance of attention between left and right space in healthy adults, and improve symptoms of spatial neglect after stroke. Here we asked whether the orienting of spatial attention to visual stimuli is affected by a different form of sensorimotor adaptation that involves physical perturbations of arm movement, rather than distortion of visual feedback. Healthy participants performed a cued discrimination task before and after they made reaching movements to a central target. A velocity‐dependent force field pushed the hand aside during each reach, and required participants to apply compensatory forces toward the opposite side. We used event‐related potentials (ERPs) to determine whether electroencephalography (EEG) responses reflecting orienting (cue‐locked N1) and disengagement (target‐locked P1) of spatial attention are affected by adaptation to force fields. After adaptation, the cue‐locked N1 was relatively larger for stimuli presented in the hemispace corresponding to the direction of compensatory hand force. P1 amplitudes evoked by invalidly cued targets presented on the opposite side were reduced. This suggests that force field adaptation boosted attentional orienting responses toward the side of hand forces, and impeded attentional disengagement from that side, mimicking previously reported effects of prism adaptation. Thus, remapping between motor commands and intended movement direction is sufficient to bias ERPs, reflecting changes in the orienting of spatial attention in the absence of visuo‐spatial distortion or visuo‐proprioceptive mismatch. Findings are relevant to theories of how sensorimotor adaptation can modulate attention, and may open new avenues for treatment of spatial neglect.     

Effects of the availability of accurate proprioceptive information on older adults' postural sway and muscle co‐contraction

During conditions of increased postural instability, older adults exhibit greater lower limb muscle co‐contraction. This response has been interpreted as a compensatory postural strategy, which may be used to increase proprioceptive information from muscle spindles or to stiffen the lower limb as a general response to minimise postural sway. The current study aimed to test these two hypotheses by investigating use of muscle co‐contraction during sensory transitions that manipulated proprioceptive input. Surface EMG was recorded from the bilateral tibialis anterior and gastrocnemius medialis muscles, in young (aged 18–30) and older adults (aged 68–80) during blind‐folded postural assessment. This commenced on a fixed platform (baseline: 2 min), followed by 3 min on a sway‐referenced platform (adaptation) and a final 3 min on a fixed platform again (reintegration). Sensory reweighting was slower in older adults, as shown by a significantly larger and longer postural sway after‐effect once a stable platform was restored. Muscle co‐contraction showed similar after‐effects, whereby older adults showed a larger increase in co‐contraction once the stable platform had been restored, compared to young adults. This co‐contraction after‐effect did not return to baseline until after 1 min. Our evidence for high muscle co‐contraction during the reintroduction of veridical proprioceptive input suggests that increased co‐contraction in older adults is not dependent on contemporaneous proprioceptive input. Rather, it is more likely that co‐contraction is a general postural strategy used to minimise postural sway, which is increased during this sensory transition. Future research should examine whether muscle co‐contraction is typically a reactive or anticipatory response.     

Neural correlates of visuomotor adjustments during scaling of human finger movements

Visually guided finger movements include online feedback of current effector position to guide target approach. This visual feedback may be scaled or otherwise distorted by unpredictable perturbations. Although adjustments to visual feedback scaling have been studied before, the underlying brain activation differences between upscaling (visual feedback larger than real movement) and downscaling (feedback smaller than real movement) are currently unknown. Brain activation differences between upscaling and downscaling might be expected because within‐trial adjustments during upscaling require corrective backwards accelerations, whereas correcting for downscaling requires forward accelerations. In this behavioural and fMRI study we investigated adjustments during up‐ and downscaling in a target‐directed finger flexion–extension task with real‐time visual feedback. We found that subjects made longer and more complete within‐trial corrections for downscaling perturbations than for upscaling perturbations. The finger task activated primary motor (M1) and somatosensory (S1) areas, premotor and parietal regions, basal ganglia, and cerebellum. General scaling effects were seen in the right pre‐supplementary motor area, dorsal anterior cingulate cortex, inferior parietal lobule, and dorsolateral prefrontal cortex. Stronger activations for down‐ than for upscaling were observed in M1, supplementary motor area (SMA), S1 and anterior cingulate cortex. We argue that these activation differences may reflect differing online correction for upscaling vs. downscaling during finger flexion‐extension.     

Alternate trains of postural muscle vibration promote cyclic body displacement in standing parkinsonian patients

Patients suffering from Parkinson's disease (PD) have difficulties with control of stance, postural changes, and walking, seemingly depending on problems in central integration of proprioceptive information. We tested the hypothesis that alternate vibration of postural muscles can induce cyclic medio‐lateral or antero‐posterior sway in PD, mimicking that accompanying body progression during walking, thereby favoring the production of locomotor tasks. In 12 standing PD patients and 11 healthy subjects, we applied trains of vibratory stimuli, bilaterally in an alternating paradigm to soleus, tibialis anterior, or paravertebral muscles. The trains of stimuli were delivered at frequencies selected to be above, near, and below the normal walking rhythm. The displacement of the center of foot pressure (CoP) was recorded. In PD, sway area during unperturbed stance was just larger than in healthy subjects; shifts in CoP in response to vibration were preserved, regardless of the vibrated muscle pair; CoP oscillations along medio‐lateral, but not antero‐posterior direction, were coupled to the vibration trains; time to initiate and terminate the postural responses was normal. PD patients correctly integrate and exploit the vibration‐induced proprioceptive inflow to produce body oscillations comparable to those occurring during walking. Vibratory stimulation can be safely and easily employed to provoke rhythmic postural changes in PD. © 2008 Movement Disorder Society     

Modulation of proprioceptive integration in the motor cortex shapes human motor learning

Sensory and motor systems interact closely during movement performance. Furthermore, proprioceptive feedback from ongoing movements provides an important input for successful learning of a new motor skill. Here, we show in humans that attention to proprioceptive input during a purely sensory task can influence subsequent learning of a novel motor task. We applied low-amplitude vibration to the abductor pollicis brevis (APB) muscle of eight healthy volunteers for 15 min while they discriminated either a small change in vibration frequency or the presence of a simultaneous weak cutaneous stimulus. Before and after the sensory attention tasks, we evaluated the following in separate experiments: (1) sensorimotor interaction in the motor cortex by testing the efficacy of proprioceptive input to reduce GABA(A)ergic intracortical inhibition using paired-pulse transcranial magnetic stimulation, and (2) how well the same subjects learned a ballistic thumb abduction task using the APB muscle. Performance of the vibration discrimination task increased the interaction of proprioceptive input with motor cortex excitability in the APB muscle, whereas performance in the cutaneous discrimination task had the opposite effect. There was a significant correlation between the integration of proprioceptive input in the motor cortex and the motor learning gain: increasing the integration of proprioceptive input from the APB increased the rate of motor learning and reduced performance variability, while decreasing proprioceptive integration had opposite effects. These findings suggest that the sensory attention tasks transiently change how proprioceptive input is integrated into the motor cortex and that these sensory changes drive subsequent learning behavior in the human motor cortex.     

High‐frequency peripheral vibration decreases completion time on a number of motor tasks

A recent theoretical account of motor control proposes that modulation of afferent information plays a role in affecting how readily we can move. Increasing the estimate of uncertainty surrounding the afferent input is a necessary step in being able to move. It has been proposed that an inability to modulate the gain of this sensory information underlies the cardinal symptoms of Parkinson's disease (PD). We aimed to test this theory by modulating the uncertainty of the proprioceptive signal using high‐frequency peripheral vibration, to determine the subsequent effect on motor performance. We investigated if this peripheral stimulus might modulate oscillatory activity over the sensorimotor cortex in order to understand the mechanism by which peripheral vibration can change motor performance. We found that 80 Hz peripheral vibration applied to the right wrist of a total of 54 healthy human participants reproducibly improved performance across four separate randomised experiments on a number of motor control tasks (nine‐hole peg task, box and block test, reaction time task and finger tapping). Improved performance on all motor tasks (except the amplitude of finger tapping) was also seen for a sample of 18PD patients ON medication. EEG data investigating the effect of vibration on oscillatory activity revealed a significant decrease in beta power (15–30 Hz) over the contralateral sensorimotor cortex at the onset and offset of 80 Hz vibration. This finding is consistent with a novel theoretical account of motor initiation, namely that modulating uncertainty of the proprioceptive afferent signal improves motor performance potentially by gating the incoming sensory signal and allowing for top‐down proprioceptive predictions.     

The predictability of frequency‐altered auditory feedback changes the weighting of feedback and feedforward input for speech motor control

Speech production requires the combined effort of a feedback control system driven by sensory feedback, and a feedforward control system driven by internal models. However, the factors that dictate the relative weighting of these feedback and feedforward control systems are unclear. In this event‐related potential (ERP) study, participants produced vocalisations while being exposed to blocks of frequency‐altered feedback (FAF) perturbations that were either predictable in magnitude (consistently either 50 or 100 cents) or unpredictable in magnitude (50‐ and 100‐cent perturbations varying randomly within each vocalisation). Vocal and P1–N1–P2 ERP responses revealed decreases in the magnitude and trial‐to‐trial variability of vocal responses, smaller N1 amplitudes, and shorter vocal, P1 and N1 response latencies following predictable FAF perturbation magnitudes. In addition, vocal response magnitudes correlated with N1 amplitudes, vocal response latencies, and P2 latencies. This pattern of results suggests that after repeated exposure to predictable FAF perturbations, the contribution of the feedforward control system increases. Examination of the presentation order of the FAF perturbations revealed smaller compensatory responses, smaller P1 and P2 amplitudes, and shorter N1 latencies when the block of predictable 100‐cent perturbations occurred prior to the block of predictable 50‐cent perturbations. These results suggest that exposure to large perturbations modulates responses to subsequent perturbations of equal or smaller size. Similarly, exposure to a 100‐cent perturbation prior to a 50‐cent perturbation within a vocalisation decreased the magnitude of vocal and N1 responses, but increased P1 and P2 latencies. Thus, exposure to a single perturbation can affect responses to subsequent perturbations.     

Instantaneous interjoint rescaling and adaptation to balance perturbation under muscular fatigue

Adaptation of automatic postural responses (APR) to balance perturbations might be thought to be impaired by muscle fatigue, given the associated proprioceptive and effector deficits. In this investigation, we aimed to evaluate the effect of muscular fatigue on APR adaptation over repetitive balance perturbations through support base backward translations. APR adaptation was evaluated in three epochs: (a) pre‐fatigue; (b) post‐fatigue, immediately following fatigue of the plantiflexor muscles through isometric contractions and (c) post‐recovery, 30 min after the end of fatiguing activity. Results showed the following: (a) Decreasing amplitudes of joints' maximum excursion over repetitive perturbations in the three fatigue‐related epochs. (b) Modulation of joints' excursion was observed in the first trial in the post‐fatigue epoch. (c) In the post‐fatigue epoch, we found interjoint rescaling, with greater amplitude of hip rotation associated with reduced amplitude of ankles' rotation. (d) Amplitudes of ankles' rotation were similar between the post‐fatigue and post‐recovery epochs. These findings lead to the conclusions that adaptation of automatic postural responses over repetitive trials was effective under focal muscular fatigue; modulation of the postural response took place in the first perturbation under fatigue, and generalization of response characteristics from post‐fatigue to post‐recovery suggests that feedforward processes in APRs generation are affected by the recent history of postural responses to stance perturbations.     

Reduced motor imagery efficiency is associated with online control difficulties in children with probable developmental coordination disorder

Recent evidence indicates that the ability to correct reaching movements in response to unexpected target changes (i.e., online control) is reduced in children with developmental coordination disorder (DCD). Recent computational modeling of human reaching suggests that these inefficiencies may result from difficulties generating and/or monitoring internal representations of movement. This study was the first to test this putative relationship empirically. We did so by investigating the degree to which the capacity to correct reaching mid-flight could be predicted by motor imagery (MI) proficiency in a sample of children with probable DCD (pDCD). Thirty-four children aged 8 to 12 years (17 children with pDCD and 17 age-matched controls) completed the hand rotation task, a well-validated measure of MI, and a double-step reaching task (DSRT), a protocol commonly adopted to infer one's capacity for correcting reaching online. As per previous research, children with pDCD demonstrated inefficiencies in their ability to generate internal action representations and correct their reaching online, demonstrated by inefficient hand rotation performance and slower correction to the reach trajectory following unexpected target perturbation during the DSRT compared to age-matched controls. Critically, hierarchical moderating regression demonstrated that even after general reaching ability was controlled for, MI efficiency was a significant predictor of reaching correction efficiency, a relationship that was constant across groups. Ours is the first study to provide direct pilot evidence in support of the view that a decreased capacity for online control of reaching typical of DCD may be associated with inefficiencies generating and/or using internal representations of action.     

Finger muscle control in children with dystonia

Background: Childhood dystonia is a disorder that involves inappropriate muscle activation during attempts at voluntary movement. Few studies have investigated the muscle activity associated with dystonia in children, and none have done so in the hands. Methods: In this study, we measured surface electromyographic activity in four intrinsic hand muscles while participants attempted to perform an isometric tracking task using one of the muscles. Results: Children with dystonia had greater tracking error with the task‐related muscle and greater overflow to non‐task muscles. Both tracking error and overflow correlated with the Barry‐Albright Dystonia scale of the respective upper limb. Overflow also decreased when participants received visual feedback of non‐task muscle activity. Dicussion: We conclude that two of the motor deficits in childhood dystonia—motor overflow and difficulties in actively controlling muscles—can be seen in the surface electromyographic activity of individual muscles during an isometric task. As expected from results in adults, overflow is an important feature of childhood dystonia. However, overflow may be at least partially dependent on an individual's level of awareness of their muscle activity. Most importantly, poor single‐muscle tracking shows that children with dystonia have deficits of individual muscle control in addition to overflow or co‐contraction. These results provide the first quantitative measures of the muscle activity associated with hand dystonia in children, and they suggest possible directions for control of dystonic symptoms. © 2011 Movement Disorder Society     

Online movement control in multiple sclerosis patients with tremor: effects of tendon vibration.

Patients with intention tremor due to multiple sclerosis (MS) exhibit an increased reliance on visual feedback in the sensorimotor control of slow goal-directed movements. In the present study, the use of proprioceptive information was investigated in MS patients with intention tremor compared to MS patients without tremor and healthy controls. Tendon vibration was applied to the wrist extensor muscles during a memory-guided slow wrist step-tracking task to investigate the use of muscle spindle afferent information in online movement control. A significant reduction of movement amplitude was induced by tendon vibration in all three groups, but the effect was found to be smaller in MS patients with tremor (28%) than in subjects without tremor (50%). Vibration also induced an increase of overall tremor amplitude in the MS tremor group; however, its effect on movement amplitude was not directly related to (changes in) tremor severity. The results suggest that the decreased online use of proprioceptive information in MS patients with tremor reflects an adaptation over time to cope with a tremor-related noisy background. Abnormalities in proprioceptive processing may explain why MS patients with tremor show an increased reliance on visual feedback for online motor control.     

Evaluating visual bias and effect of proprioceptive feedback in unilateral neglect

We propose modified versions of the line-bisection task for assessing visual bias and effect of proprioceptive feedback in unilateral neglect; that is, the verbal line-bisection (VLB) task and the VLB plus pointing task. The VLB task requires only a verbal response to assess pure visual bias. Conversely, the VLB plus pointing task requires both a verbal response and pointing to assess visual and proprioceptive bias. Ten patients with unilateral neglect were administered these tasks on a computer using presentation software. Eight participants showed obvious rightward deviation in the VLB task, and were thus classified as patients with visual neglect. Four participants showed significantly greater deviation in the VLB plus pointing task than in the VLB task, and were thus classified as patients with proprioceptive bias. Visual bias and effect of proprioceptive feedback in unilateral neglect were successfully assessed by these tasks.     

Dependence of cerebellar tremor on proprioceptive but not visual feedback

We studied the influence of proprioceptive and visual feedback on cerebellar tremor which occurred after arm perturbations and after voluntary elbow flexions. Cerebellar tremor was produced in monkeys by reversibly cooling through two probes implanted lateral and medial to the dentate nucleus. Cerebellar tremor was synchronized in different trials to torque pulse onset and to the end, but not the start, of voluntary movements. Addition of loads to the handle held by the monkey (increases in spring stiffness, viscosity, constant torque, and inertial load) changed the amplitude and frequency of tremor that follows arm perturbations or voluntary movements in the same way. In both situations EMG activity in each cycle of tremor followed stretch of its own muscle and attained a peak near peak velocity irrespective of the mechanical load. Removal of visual feedback did not alter the characteristics of the tremor or the associated EMG activity. We concluded that cerebellar intention tremor, which occurs when attempting to hold the arm in an intended position, is driven by stretch-evoked peripheral feedback and not by voluntary corrections based on vision.     

A task-dependent effect of memory and hand-target on proprioceptive localization

We examine whether the task goal affects the accuracy and precision with which participants can localize an unseen hand. Proprioceptive localization was measured using three different tasks: two goal-directed movement tasks (reaching to and reproducing final hand-target location) and a perceptual estimation task in which participants judged the location of the hand-target relative to visual references. We also assessed whether proprioceptive localization in these different tasks is affected by localization from memory, the hand-target being localized (left or right) or the movement path of the proprioceptive target (9 paths, derived from combinations of starting and final hand-target positions). We found that participants were less precise when reaching from memory, but not when reproducing or estimating remembered final hand-target location. Participants also misperceived the felt location of their hands, judging their left hand to be more leftward and their right hand to be more rightward when reaching to and when estimating final hand-target location, but not when reproducing hand-target location. The movement path of the proprioceptive target did not affect localization, regardless of the task goal. Overall, localization seems poorer when proprioception is used to guide a reach with the opposite hand, particularly from memory, and best when merely reproducing the proprioceptive target site. This may have an important application in neuro-rehabilitation, whereby one task may better establish or re-establish important or failing sensory connections.     

Task-relevant selective modulation of somatosensory afferent paths from the lower limb

Leg movement attenuates initial somatosensory evoked potentials (SEPS) from both cutaneous and muscle afferent origin. To date, as different sensory inputs become relevant for task performance, selective facilitation from such movement-related gating influences has not been shown. We hypothesized that initial SEP amplitudes from cutaneous (sural nerve, SN) and muscle afferent (tibial nerve, TN) sources are dependent on the relevance of the specific afferent information to task performance. SEPs were obtained at rest and during three movement conditions. In each movement condition, the left foot was passively moved episodically and additional cutaneous 'codes' of sensory information were applied to the dorsum of the left foot. Subjects were instructed to: simply relax (passive), or to make a response following the cessation of movement, dependent either on the cutaneous code (cutaneous task), or the passive movement trajectory of the left foot (position task). Passive movement, with no required subsequent response, attenuated initial TN and SN SEPs to approximately 40% of that at rest (p < 0.05). Versus passive movement, when cutaneous inputs provided the relevant cue for the task, mean SN SEPs significantly increased (p < 0.05), and when the proprioceptive inputs provided the relevant cue for the task, mean TN SEPs significantly increased (p < 0.05). We conclude that specific relevancy of sensory information selectively facilitates somatosensory paths from movement-related attenuation.     

Directional communication during movement execution interferes with tremor in Parkinson's disease

Background : Both the cerebello‐thalamo‐cortical circuit and the basal ganglia/cortical motor loop have been postulated to be generators of tremor in PD. The recent suggestion that the basal ganglia trigger tremor episodes and the cerebello‐thalamo‐cortical circuitry modulates tremor amplitude combines both competing hypotheses. However, the role of the STN in tremor generation and the impact of proprioceptive feedback on tremor suppression during voluntary movements have not been considered in this model yet. Objectives : The objective of this study was to evaluate the role of the STN and proprioceptive feedback in PD tremor generation during movement execution. Methods: Local‐field potentials of the STN as well as electromyographical and electroencephalographical rhythms were recorded in tremor‐dominant and nontremor PD patients while performing voluntary movements of the contralateral hand during DBS surgery. Effective connectivity between these electrophysiological signals were analyzed and compared to electromyographical tremor activity. Results: There was an intensified information flow between the STN and the muscle in the tremor frequencies (5‐8 Hz) for tremor‐dominant, in comparison to nontremor, patients. In both subtypes, active movement was associated with an increase of afferent interaction between the muscle and the cortex in the β‐ and γ‐frequencies. The γ‐frequency (30‐40 Hz) of this communication between muscle and cortex correlated inversely with electromyographical tremor activity. Conclusions : Our results indicate an involvement of the STN in propagation of tremor‐related activity to the muscle. Furthermore, we provide evidence that increased proprioceptive information flow during voluntary movement interferes with central tremor generation. © 2018 International Parkinson and Movement Disorder Society     

Cervical dystonia affects aimed movements of nondystonic segments

Patients with focal dystonia exhibit proprioception abnormalities that can lead to kinematic deficits. Proprioceptive abnormalities are present in both symptomatic and asymptomatic body parts of dystonic patients. To ascertain whether in patients with idiopathic cervical dystonia (CD) movements performed with nondystonic segments display kinematic abnormalities, we studied trajectory formation of out and back arm reaching movements in 10 patients with CD (before and 3 weeks after treatment with Botulinum toxin) and in 10 age‐matched controls. Before treatment, patients with CD showed significant trajectory abnormalities when compared with normal controls. Patients' trajectories were more curved with asymmetrical temporal velocity profiles as well as increased hand path areas, and had longer reversal lags between the out and back segments. Treatment with botulinum toxin improved all the kinematic parameters. These results suggest that in patients with CD, movements performed with nondystonic segments are abnormal. The kinematic abnormalities are likely to derive from long‐standing defective integration of the proprioceptive input, which, in turn, causes general changes in the internal models of limb dynamics. It is plausible that treatment with botulinum toxin partially restores proprioceptive processing and thus, such internal models. © 2009 Movement Disorder Society     

Feedback and feedforward adaptation to visuomotor delay during reaching and slicing movements

It has been suggested that the brain and in particular the cerebellum and motor cortex adapt to represent the environment during reaching movements under various visuomotor perturbations. It is well known that significant delay is present in neural conductance and processing; however, the possible representation of delay and adaptation to delayed visual feedback has been largely overlooked. Here we investigated the control of reaching movements in human subjects during an imposed visuomotor delay in a virtual reality environment. In the first experiment, when visual feedback was unexpectedly delayed, the hand movement overshot the end‐point target, indicating a vision‐based feedback control. Over the ensuing trials, movements gradually adapted and became accurate. When the delay was removed unexpectedly, movements systematically undershot the target, demonstrating that adaptation occurred within the vision‐based feedback control mechanism. In a second experiment designed to broaden our understanding of the underlying mechanisms, we revealed similar after‐effects for rhythmic reversal (out‐and‐back) movements. We present a computational model accounting for these results based on two adapted forward models, each tuned for a specific modality delay (proprioception or vision), and a third feedforward controller. The computational model, along with the experimental results, refutes delay representation in a pure forward vision‐based predictor and suggests that adaptation occurred in the forward vision‐based predictor, and concurrently in the state‐based feedforward controller. Understanding how the brain compensates for conductance and processing delays is essential for understanding certain impairments concerning these neural delays as well as for the development of brain–machine interfaces.     

Tonic vibration reflex and muscle afferent block in writer's cramp

Patients with focal dystonia take advantage of certain cutaneous or proprioceptive sensory inputs to alleviate their symptoms (“sensory trick”). We examined the effects of increasing muscle spindle activity by the tonic vibration reflex maneuver and Decemberreasing it by intramuscular injection of lidocaine. The vibration was applied to the palm or the tendon of forearm muscles in 15 patients with writer's cramp and 15 age-matched normal subjects. In 11 patients, the vibration induced dystonic postures or movements typical of those seen during writing. Normal subjects showed either no response to the vibration or a gradually developing tonic vibration reflex only in the wrist and finger flexors, which produced visible movements with a significantly longer latency (12.5 ± 6.7 seconds [mean ± standard deviation]) than what was observed in the patients (2.7 ± 2.5 seconds, p < 0.0001). Local injection of lidocaine (0.5%, 5–40 ml/muscle) attenuated the tendon reflex with relatively little effect on the M response. Injection into muscles with increased activity produced Marchked reduction of dystonic movements and significant clinical improvement in 13 patients, whereas injection into the other muscles had no effect. The clinical benefit lasted for 1 to 24 hours after injection. In 13 patients who had additional injections of 10% ethanol, which blocks sodium channels for a longer period than does lidocaine, the duration of action was prolonged to 5 to 21 days. These findings suggest that muscles causing dystonic movements have abnormal sensitivities to vibration at rest and that muscle afferents may play a pivotal role in producing dystonic movements. Local injection of lidocaine or ethanol reduces the effectiveness of muscle spindle afferent and may provide a means of treating patients with writer's cramp without causing unwanted weakness.