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1.  Spatial and Temporal Asymmetries in Gait Predict Split-Belt Adaptation Behavior in Stroke 
Step asymmetries during gait in persons after stroke can occur in temporal or spatial domains. Prior studies have shown that split-belt locomotor adaptation can temporarily mitigate these asymmetries.
We investigated whether baseline gait asymmetries affected how patients adapt and store new walking patterns.
Subjects with stroke and age-matched controls were studied walking at a 2:1 speed ratio on the split-belt during adaptation and assessed for retention of the learned pattern (the after-effect) with both belts at the same speed.
Those with stroke adapted more slowly (P < .0001), though just as much as healthy older adults. During split-belt walking, the participants with stroke adapted toward their baseline asymmetry (eg, F = 14.02, P < .01 for step symmetry), regardless of whether the subsequent after-effects improved or worsened their baseline step asymmetries. No correlation was found between baseline spatial and temporal measures of asymmetry (P = .38). Last, the initial spatial and temporal asymmetries predicted after-effects independently of one another. The after-effects in the spatial domain (ie, center of oscillation difference) are only predicted by center of oscillation difference baseline (F = 15.3, P = .001), while all other parameters were nonsignificant (all Ps > .17). Temporal coordination (ie, phasing) after-effects showed a significant effect only from phasing baseline (F = 26.92, P < .001, all others P > .33).
This work demonstrates that stroke patients adapt toward their baseline temporal and spatial asymmetries of walking independently of one another. We define how a given split-belt training session would affect asymmetries in these domains, which must be considered when developing rehabilitation interventions for stroke patients.
PMCID: PMC4336782  PMID: 24243917
locomotor rehabilitation; split-belt treadmill; motor adaptation; poststroke; kinematics; walking
2.  ‘Motor cognition’ – what is it and is the cerebellum involved? 
Cerebellum (London, England)  2007;6(3):232-236.
Motor cognition encompasses how we understand our own movement, and how movement helps us to understand the world. Here, the role of the cerebellum is discussed in two processes that could be considered aspects of motor cognition: predicting movement outcomes and understanding the meaning of movements. Recent behavioral, anatomical, and neurophysiological findings related to these processes are discussed. There are data to support a cerebellar role in predicting movement outcomes, which could be used both for motor control and for distinguishing sensory inputs due to our own movements from external influences. The data for a cerebellar role in understanding the meaning of movement are mixed, although anatomical findings suggest that it probably has some influence that bears further study.
PMCID: PMC4096570  PMID: 17786819
movement; prediction; autism; schizophrenia
3.  Predictive Modeling by the Cerebellum Improves Proprioception 
The Journal of Neuroscience  2013;33(36):14301-14306.
Because sensation is delayed, real-time movement control requires not just sensing, but also predicting limb position, a function hypothesized for the cerebellum. Such cerebellar predictions could contribute to perception of limb position (i.e., proprioception), particularly when a person actively moves the limb. Here we show that human cerebellar patients have proprioceptive deficits compared with controls during active movement, but not when the arm is moved passively. Furthermore, when healthy subjects move in a force field with unpredictable dynamics, they have active proprioceptive deficits similar to cerebellar patients. Therefore, muscle activity alone is likely insufficient to enhance proprioception and predictability (i.e., an internal model of the body and environment) is important for active movement to benefit proprioception. We conclude that cerebellar patients have an active proprioceptive deficit consistent with disrupted movement prediction rather than an inability to generally enhance peripheral proprioceptive signals during action and suggest that active proprioceptive deficits should be considered a fundamental cerebellar impairment of clinical importance.
PMCID: PMC3761044  PMID: 24005283
4.  Cerebellar involvement in motor but not sensory adaptation 
Neuropsychologia  2012;50(8):1766-1775.
Predictable sensorimotor perturbations can lead to cerebellum-dependent adaptation—i.e., recalibration of the relationship between sensory input and motor output. Here we asked if the cerebellum is also needed to recalibrate the relationship between two sensory modalities, vision and proprioception. We studied how people with and without cerebellar damage use visual and proprioceptive signals to estimate their hand’s position when the sensory estimates disagree. Theoretically, the brain may resolve the discrepancy by recalibrating the relationship between estimates (sensory realignment). Alternatively, the misalignment may be dealt with by relying less on one sensory estimate and more on the other (a weighting strategy). To address this question, we studied subjects with cerebellar damage and healthy controls as they performed a series of tasks. The first was a prism adaptation task that involves motor adaptation to compensate for a visual perturbation and is known to require the cerebellum. As expected, people with cerebellar damage were impaired relative to controls. The same subjects then performed two experiments in which they reached to visual and proprioceptive targets while a visuoproprioceptive misalignment was gradually imposed. Surprisingly, cerebellar patients performed as well as controls when the task invoked only sensory realignment, but were impaired relative to controls when motor adaptation was also possible. Additionally, individuals with cerebellar damage were able to use a weighting strategy similarly to controls. These results demonstrate that, unlike motor adaptation, sensory realignment and weighting are not cerebellum-dependent.
PMCID: PMC3389289  PMID: 22554563
motor adaptation; sensory adaptation; sensorimotor integration; reaching; cerebellum
5.  Longitudinal Tracking of Gait and Balance Impairments in Cerebellar Disease 
Cerebellar damage typically results in ataxia and can be caused by stroke, tumor or one of many forms of degenerative disease. Since few pharmacological options are available, most treatments rely heavily on rehabilitation therapy. Little data exist on methods for tracking the progression of ataxia, which is critical for assessing the efficacy of current and newly developing treatments. Here, we tracked the severity of ataxia, with a particular emphasis on gait and balance dysfunction, in a group of individuals with cerebellar damage using the International Cooperative Ataxia Rating Scale (ICARS) and several instrumented laboratory measures of gait and balance impairments over one year. We found that the ICARS was able to distinguish between subjects with static lesions and those with degenerative disorders, was sensitive to increases in ataxia severity occurring over one year, and correlated well with specific instrumented measures of gait in persons with cerebellar degeneration. These results suggest the ICARS is a valuable tool for clinicians and investigators to document and track long-term changes in gait and balance performance in individuals with cerebellar degenerative disorders.
PMCID: PMC2939173  PMID: 20544808
ICARS; cerebellum; walking; ataxia; clinical assessment; sensitivity
6.  Younger is not always better: Development of locomotor adaptation from childhood to adulthood 
New walking patterns can be learned over short timescales (i.e. adapted in minutes) using a split-belt treadmill that controls the speed of each leg independently. This leads to storage of a modified spatial and temporal motor pattern that is expressed as an aftereffect in regular walking conditions. Since split-belt walking is a novel task for adults and children alike, we used it to investigate how motor adaptation matures during human development. We also asked if the immature pattern resembles that of people with cerebellar dysfunction, since we know that this adaptation depends on cerebellar integrity. Healthy children (3–18 years) and adults, and individuals with cerebellar damage were adapted while walking on split-belts (1:2 speed ratio). Adaptation and de-adaptation rates were quantified separately for temporal and spatial parameters. All healthy children and adults tested could learn the new timing at the same rate, and showed significant aftereffects. However, children younger than six years were unable to learn the new spatial coordination. Further, children as old as age 11 showed slower rates of adaptation and de-adaptation of spatial parameters of walking. Young children showed patterns similar to cerebellar patients, with greater deficits in spatial versus temporal adaptation. Thus, even though walking is a well-practiced, refined motor skill by late childhood (i.e. 11 years of age), the processes underlying learning new spatial relationships between the legs are still developing. The maturation of locomotor adaptation follows at least two time courses, which we propose is determined by the developmental state of the cerebellum.
PMCID: PMC3084584  PMID: 21414926
Locomotion; adaptation; motor learning; development; interlimb coordination; split-belt treadmill
7.  Seeing is believing: effects of visual contextual cues on learning and transfer of locomotor adaptation 
Devices such as robots or treadmills are often used to drive motor learning because they can create novel physical environments. However, the learning (i.e., adaptation) acquired on these devices only partially generalizes to natural movements. What determines the specificity of motor learning, and can this be reliably made more general? Here we investigated the effect of visual cues on the specificity of split-belt walking adaptation. We systematically removed vision to eliminate the visual-proprioceptive mismatch that is a salient cue specific to treadmills: vision indicates that we are not moving while leg proprioception indicates that we are. We evaluated the adaptation of temporal and spatial features of gait (i.e., timing and location of foot landing), their transfer to walking over ground, and washout of adaptation when subjects returned to the treadmill. Removing vision during both training (i.e., on the treadmill) and testing (i.e., over ground) strongly improved the transfer of treadmill adaptation to natural walking. Removing vision only during training increased transfer of temporal adaptation, whereas removing vision only during testing increased the transfer of spatial adaptation. This dissociation reveals differences in adaptive mechanisms for temporal and spatial features of walking. Finally training without vision increased the amount that was learned and was linked to the variability in the behavior during adaptation. In conclusion, contextual cues can be manipulated to modulate the magnitude, transfer, and washout of device-induced learning in humans. These results bring us closer to our ultimate goal of developing rehabilitation strategies that improve movements beyond the clinical setting.
PMCID: PMC3025449  PMID: 21159971
human; generalization; locomotion; motor control; kinematics; motor learning; vision
8.  Understanding sensorimotor adaptation and learning for rehabilitation 
Current opinion in neurology  2008;21(6):628-633.
Purpose of review
Understanding the behavioral mechanisms of sensorimotor adaptation and learning is essential for designing rational rehabilitation interventions.
Recent findings
Adaptation is the trial-and-error process of adjusting movement to new demands and is now thought to be more than a simple error cancellation process. Instead, it may calibrate the brain’s prediction of how the body will move and takes into account costs associated with the new task demand. Damage of the cerebellum systematically disrupts adaptation, but damage to other brain regions often does not. Adapting to perturbations driven by a device like a robot or a treadmill leads to only partial generalization to unconstrained ‘real-world’ movements. Repeated adaptation can lead to learning a new motor calibration, but process of consolidation of this type of learning is less understood in patients.
Adaptation is inherently important for rehabilitation by making movement flexible, but can also be used to ascertain whether some patients can generate a more normal motor pattern. Repeated adaptation can lead to learning of a new, more permanent motor calibration. Though less understood, this type of learning is likely to be an important method for making long-term improvements in patients’ movement patterns.
PMCID: PMC2954436  PMID: 18989103
memory; motor; reaching; stroke; walking
9.  Prevalence of ataxia in children 
Neurology  2014;82(1):80-89.
To estimate the prevalence of childhood ataxia resulting from both genetic and acquired causes.
A systematic review was conducted following the PRISMA (Preferred Reporting Items for Systematic Reviews and Meta-analyses) statement. Five databases were searched for articles reporting a frequency measure (e.g., prevalence, incidence) of ataxia in children. Included articles were first grouped according to the World Health Organization (WHO) regions and subsequently classified according to etiology (genetic, acquired, or mixed). Each article was assessed for its risk of bias on the domains of sampling, measurement, and analysis. Incidence values were converted to prevalence estimates whenever possible. European prevalence estimates for different etiologies of ataxia were summed to gauge the overall prevalence of childhood ataxia.
One hundred fifteen articles were included in the review. More than 50% of the data originated from the Europe WHO region. Data from this region also showed the least susceptibility to bias. Little data were available for Africa and Southeast Asia. The prevalence of acquired ataxias was found to vary more greatly across regions than the genetic ataxias. Ataxic cerebral palsy was found to be a significant contributor to the overall prevalence of childhood ataxia across WHO regions. The prevalence of childhood ataxias in Europe was estimated to be ∼26/100,000 children and likely reflects a minimum prevalence worldwide.
The findings show that ataxia is a common childhood motor disorder with a higher prevalence than previously assumed. More research concerning the epidemiology, assessment, and treatment of childhood ataxia is warranted.
PMCID: PMC3873624  PMID: 24285620
10.  The effects of task demands on bimanual skill acquisition 
Bimanual coordination is essential for everyday activities. It is thought that different degrees of demands may affect learning of new bimanual patterns. One demand is at the level of performance and involves breaking the tendency to produce mirror-symmetric movements. A second is at a perceptual level and involves controlling each hand to separate (i.e., split) goals. A third demand involves switching between different task contexts (e.g., a different uni- or bimanual task), instead of continuously practicing one task repeatedly. Here, we studied the effect of these task demands on motor planning (reaction time) and execution (error) while subjects learned a novel bimanual isometric pinch force task. In Experiment 1, subjects continuously practiced in one of the two extremes of the following bimanual conditions: (1) symmetric force demands and a perceptually unified target for each hand or (2) asymmetric force demands and perceptually split targets. Subjects performing in the asymmetric condition showed some interference between hands, but all subjects, regardless of group, could learn the isometric pinch force task similarly. In Experiment 2, subjects practiced these and two other conditions, but in a paradigm where practice was briefly interrupted by the performance of either a unimanual or a different bimanual condition. Reaction times were longer and errors were larger well after the interruption when the main movement to be learned required asymmetric forces. There was no effect when the main movement required symmetric forces. These findings demonstrate two main points. First, people can learn bimanual tasks with very different demands on the same timescale if they are not interrupted. Second, interruption during learning can negatively impact both planning and execution and this depends on the demands of the bimanual task to be learned. This information will be important for training patient populations, who may be more susceptible to increased task demands.
PMCID: PMC3615041  PMID: 23392473
Bimanual; Skill learning; Motor control; Task demands
11.  Prior Experience but Not Size of Error Improves Motor Learning on the Split-Belt Treadmill in Young Children 
PLoS ONE  2014;9(3):e93349.
Children can modify learned motor skills, such as walking, to adapt to new environments. Movement errors in these new situations drive the learning. We used split-belt walking to determine whether size of the error affects the degree of learning. Twenty-two children (aged 2–5 y) walked on the split-belt treadmill on two separate days spaced 1 week apart. Twenty-eight adults served as controls. On Day 1, children experienced an abrupt change in belt speeds (from 1∶1 to 2∶1 differential) resulting in large errors, or a gradual change (same change in speed over 12–15 min), resulting in small errors. Learning was measured by the size of the aftereffect upon return to a 1∶1 differential. On Day 2 (1 week later), the leg on the fast belt was reversed, as was the method of introducing the speed differential. We found that the error size did not affect learning. Unexpectedly, learning was greater on Day 2 compared to Day 1, especially for children under 4 y of age, despite the fact that the task was opposite to that of Day 1, and did not influence learning in adults. Hence, 11 additional children under 4 y of age were tested with belts running at the same speed on Day 1, and with a 2∶1 speed differential (abrupt introduction) on Day 2. Surprisingly, learning was again greater on Day 2. We conclude that size of error during split-belt walking does not affect learning, but experience on a treadmill does, especially for younger children.
PMCID: PMC3968172  PMID: 24675816
12.  Repeated Split-BeltTreadmill Training Improves Poststroke Step Length Asymmetry 
Background and objective
Previous studies suggest that error augmentation may be used as a strategy to achieve longer-term changes in gait deficits after stroke. The purpose of this study was to determine whether longer-term improvements in step length asymmetry could be achieved with repeated split-belt treadmill walking practice using an error augmentation strategy.
13 persons with chronic stroke (>6 months) participated in testing: (1) prior to 12 sessions of split-belt treadmill training, (2) after the training, and (3) in follow-up testing at 1 and 3 months. Step length asymmetry was the target of training, so belt speeds were set to augment step length asymmetry such that aftereffects resulted in reduced step length asymmetry during overground walking practice. Each individual was classified as a “responder” or “nonresponder” based on whether their reduction in step length asymmetry exceeded day-to-day variability.
For the group and for the responders (7 individuals), step length asymmetry improved from baseline to posttesting (P < .05) through an increased step length on both legs but a relatively larger change on the shorter step side (P < .05). Other parameters that were not targeted (eg, stance time asymmetry) did not change over the intervention.
This study demonstrates that short-term adaptations can be capitalized on through repetitive practice and can lead to longer-term improvements in gait deficits poststroke. The error augmentation strategy, which promotes stride-by-stride adjustment to reduce asymmetry and results in improved asymmetry during overground walking practice, appears to be critical for obtaining the improvements observed.
PMCID: PMC3738184  PMID: 23392918
stroke; gait; motor learning
13.  Locomotor adaptation 
Progress in brain research  2011;191:65-74.
Motor learning is an essential part of human behavior, but poorly understood in the context of walking control. Here, we discuss our recent work on locomotor adaptation, which is an error driven motor learning process used to alter spatiotemporal elements of walking. Locomotor adaptation can be induced using a split-belt treadmill that controls the speed of each leg independently. Practicing split-belt walking changes the coordination between the legs, resulting in storage of a new walking pattern. Here, we review findings from this experimental paradigm regarding the learning and generalization of locomotor adaptation. First, we discuss how split-belt walking adaptation develops slowly throughout childhood and adolescence. Second, we demonstrate that conscious effort to change the walking pattern during split-belt training can speed up adaptation but worsens retention. In contrast, distraction (i.e., performing a dual task) during training slows adaptation but improves retention. Finally, we show the walking pattern acquired on the split-belt treadmill generalizes to natural walking when vision is removed. This suggests that treadmill learning can be generalized to different contexts if visual cues specific to the treadmill are removed. These findings allow us to highlight the many future questions that will need to be answered in order to develop more rational methods of rehabilitation for walking deficits.
PMCID: PMC3738197  PMID: 21741544
locomotion; motor learning; adaptation; generalization of learning; rehabilitation
14.  Dynamic modulation of cerebellar excitability for abrupt, but not gradual visuomotor adaptation 
The cerebellum is critically important for error driven adaptive motor learning, as evidenced by the fact that cerebellar patients do not adapt well to sudden predictable perturbations. However, recent work has shown that cerebellar patients adapt much better if the perturbation is gradually introduced. Here we explore physiological mechanisms that underlie this distinction between abrupt and gradual motor adaptation in humans. We used Transcranial Magnetic Stimulation (TMS) to evaluate whether neural mechanisms within the cerebellum contribute to either process during a visuomotor reach adaptation. When a visuomotor rotation was introduced abruptly, cerebellar excitability changed early in learning, and approached baseline levels near the end of the adaptation block. However, we observed no modulation of cerebellar excitability when we presented the visuomotor rotation gradually during learning. Similarly, we did not observe cerebellar modulation during trial-by-trial adaptation to random visuomotor displacements or during reaches without perturbations. This suggests that the cerebellum is most active during the early-phases of adaptation when large perturbations are successfully compensated.
PMCID: PMC3435878  PMID: 22915105
Motor Learning; M1; Error; Reaching; Stimulation
15.  No Proprioceptive Deficits in Autism Despite Movement-Related Sensory and Execution Impairments 
Autism spectrum disorder (ASD) often involves sensory and motor problems, yet the proprioceptive sense of limb position has not been directly assessed. We used three tasks to assess proprioception in adolescents with ASD who had motor and sensory perceptual abnormalities, and compared them to age- and IQ-matched controls. Results showed no group differences in proprioceptive accuracy or precision during active or passive tasks. Both groups showed (a) biases in elbow angle accuracy that varied with joint position, (b) improved elbow angle precision for active versus passive tasks, and (c) improved precision for a fingertip versus elbow angle estimation task. Thus, a primary proprioceptive deficit may not contribute to sensorimotor deficits in ASD. Abnormalities may arise at later sensory processing stages.
PMCID: PMC3118271  PMID: 21165765
Proprioception; Motor control; Sensory processing
16.  Moving, sensing and learning with cerebellar damage 
Current opinion in neurobiology  2011;21(4):596-601.
The cerebellum is a subcortical brain structure that is essential for learning and controlling movement. Recent work shows that the cerebellum also plays a role in certain perceptual abilities, beyond what would be expected secondary to poor movement control. This review covers these and other recent advances, focusing on how cerebellar damage affects human abilities ranging from sensory perception to movement control and motor learning.
PMCID: PMC3177958  PMID: 21733673
17.  Human Locomotor Adaptive Learning Is Proportional to Depression of Cerebellar Excitability 
Cerebral Cortex (New York, NY)  2011;21(8):1901-1909.
Human locomotor adaptive learning is thought to involve the cerebellum, but the neurophysiological mechanisms underlying this process are not known. While animal research has pointed to depressive modulation of cerebellar outputs, a direct correlation between adaptive learning and cerebellar depression has never been demonstrated. Here, we used transcranial magnetic stimulation to assess excitability changes occurring in the cerebellum and primary motor cortex (M1) after individuals learned a new locomotor pattern on a split-belt treadmill. To control for potential changes associated to task performance complexity, the same group of subjects was also assessed after performing 2 other locomotor tasks that did not elicit learning. We found that only adaptive learning resulted in reduction of cerebellar inhibition. This effect was strongly correlated with the magnitude of learning (r = 0.78). In contrast, M1 excitability changes were not specific to learning but rather occurred in association with task complexity performance. Our results demonstrate that locomotor adaptive learning in humans is proportional to cerebellar excitability depression. This finding supports the theory that adaptive learning is mediated, at least in part, by long-term depression in Purkinje cells. This knowledge opens the opportunity to target cerebellar processes with noninvasive brain stimulation to enhance motor learning.
PMCID: PMC3202738  PMID: 21239392
adaptation; cerebellum; locomotion; rehabilitation; TMS
Even after rehabilitation, many individuals with stroke have residual gait deviations and limitations in functional walking. Applying the principles of motor adaptation through a split-belt treadmill walking paradigm can lead to short-term improvements in step length asymmetry after stroke. The focus of this case study was to determine whether it is possible to capitalize on these improvements for long-term gain.
Case Description
The participant was a 36-year-old female who was 1.6 years post-stroke. She had a slow walking speed and multiple specific gait deviations, including step length asymmetry.
The participant walked on the split-belt treadmill 3 days/week for four weeks, with the paretic leg on slow belt. The goal was 30 minutes of split-belt treadmill walking each day, followed by overground walking practice to reinforce improvements in step length symmetry.
With training, step length asymmetry decreased from 21% to 9%, and decreased further to 7% asymmetry 1 month after training. Self-selected walking speed increased from 0.71 m/s to 0.81 m/s after training and 0.86 m/s 1 month later. Percent recovery, measured by the Stroke Impact Scale, increased from 40% to 50% post-training and to 60% 1 month later.
Improvements in step length symmetry were observed following training and these improvements were maintained 1 month later. Concomitant changes in clinical measures were also observed, although these were relatively modest. The outcomes for this participant are encouraging given the relatively small dose of training. They suggest that short-term adaptation can be capitalized on through repetitive practice and can lead to longer-term improvements after stroke.
PMCID: PMC3394680  PMID: 21084921
19.  Motor adaptation training for faster re-learning 
Adaptation is an error-driven motor learning process that can account for predictable changes in the environment (e.g. walking on ice) or in ourselves (e.g. injury). Our ability to recall and build upon adapted motor patterns across days is essential to this learning process. We investigated how different training paradigms affect the day-to-day memory of an adapted walking pattern. Healthy human adults walked on a split-belt treadmill, and returned the following day to assess recall, re-learning rate, and performance. In the first experiment, one group adapted and de-adapted (i.e. washed-out the learning) several times on day one to practice the initial stage of learning where errors are large; another group adapted only one time and then practiced in the adapted (“learned”) state where errors were small. On day two, they performed washout trials before re-adapting. The group that repeatedly practiced the initial portion of adaptation where errors are large showed the fastest re-learning on the second day. In fact, the memory was nearly as strong as that of a third group that was left overnight in the adapted state and was not washed-out prior to re-exposure on the second day. This demonstrates that alternating exposures to early adaptation and washout can enhance re-adaptation. In the second experiment, we tested whether the opposite split-belt pattern interferes with day two re-learning. Surprisingly, it did not, and instead was similar to practicing in the adapted state. These results show that the structure of the initial phase of learning influences the ease of motor re-learning.
PMCID: PMC3209529  PMID: 22016547
motor control; gait; structure of practice; motor learning; locomotor adaptation
20.  Perceptual reasoning predicts handwriting impairments in adolescents with autism 
Neurology  2010;75(20):1825-1829.
We have previously shown that children with autism spectrum disorder (ASD) have specific handwriting deficits consisting of poor form, and that these deficits are predicted by their motor abilities. It is not known whether the same handwriting impairments persist into adolescence and whether they remain linked to motor deficits.
A case-control study of handwriting samples from adolescents with and without ASD was performed using the Minnesota Handwriting Assessment. Samples were scored on an individual letter basis in 5 categories: legibility, form, alignment, size, and spacing. Subjects were also administered an intelligence test and the Physical and Neurological Examination for Subtle (Motor) Signs (PANESS).
We found that adolescents with ASD, like children, show overall worse performance on a handwriting task than do age- and intelligence-matched controls. Also comparable to children, adolescents with ASD showed motor impairments relative to controls. However, adolescents with ASD differ from children in that Perceptual Reasoning Indices were significantly predictive of handwriting performance whereas measures of motor skills were not.
Like children with ASD, adolescents with ASD have poor handwriting quality relative to controls. Despite still demonstrating motor impairments, in adolescents perceptual reasoning is the main predictor of handwriting performance, perhaps reflecting subjects' varied abilities to learn strategies to compensate for their motor impairments.
= autism spectrum disorder;
= Diagnostic and Statistical Manual of Mental Disorders, 4th edition;
= Physical and Neurological Examination for Subtle (Motor) Signs;
= Perceptual Reasoning Index;
= Wechsler Abbreviated Scale of Intelligence;
= Wechsler Intelligence Scale for Children IV.
PMCID: PMC2995383  PMID: 21079184
21.  Sensorimotor function and sensorimotor tracts after hemispherectomy 
Neuropsychologia  2009;48(5):1192-1199.
Hemispherectomy is currently the only effective treatment for relieving constant seizures in children with severe or progressive unilateral cortical disease. Although early hemispherectomy has been advocated to avoid general dysfunction due to continued seizures, it remains unclear whether age at surgery affects specific sensorimotor functions. Little is know about the anatomical status of sensorimotor pathways after hemispherectomy and how it might relate to sensorimotor function. Here we measured motor function and sensory thresholds of the upper and lower limbs in 12 hemispherectomized patients. Diffusion tensor imaging (DTI) was used to determine status of brainstem corticospinal tracts and medial lemniscus. Hemispherectomy subjects showed remarkable recovery in both sensory and motor function. Many patients showed normal sensory vibration thresholds. Within the smaller Rasmussen’s subgroup, we saw a relationship between age at surgery and sensorimotor function recovery (i.e. earlier was better). Anatomically, we found marked asymmetry in brainstem corticospinal tracts but preserved symmetry in the medial lemniscus, which may relate to robust sensory recovery. Age at surgery predicted anatomical status of brainstem sensorimotor tracts. In sum, we found that age at surgery influences anatomical changes in brainstem motor pathways, and may also relate to sensorimotor recovery patterns.
PMCID: PMC2847436  PMID: 20018199
diffusion tensor imaging; corticospinal tract; dorsal column medial lemniscal tract; vibration sensation; Fugl-Meyer assessment
22.  Children with autism show specific handwriting impairments 
Neurology  2009;73(19):1532-1537.
Handwriting skills, which are crucial for success in school, communication, and building children’s self-esteem, have been observed to be poor in individuals with autism. Little information exists on the handwriting of children with autism, without delineation of specific features that can contribute to impairments. As a result, the specific aspects of handwriting in which individuals with autism demonstrate difficulty remain unknown.
A case-control study of handwriting samples from children with and without autism spectrum disorders (ASD) was performed using the Minnesota Handwriting Assessment. Samples were scored on an individual letter basis in 5 categories: legibility, form, alignment, size, and spacing. Subjects were also tested on the Wechsler Intelligence Scale for Children–IV and the Physical and Neurological Examination for Subtle (Motor) Signs.
We found that children with ASD do indeed show overall worse performance on a handwriting task than do age- and intelligence-matched controls. More specifically, children with ASD show worse quality of forming letters but do not show differences in their ability to correctly size, align, and space their letters. Within the ASD group, motor skills were significantly predictive of handwriting performance, whereas age, gender, IQ, and visuospatial abilities were not.
We addressed how different elements of handwriting contribute to impairments observed in children with autism. Our results suggest that training targeting letter formation, in combination with general training of fine motor control, may be the best direction for improving handwriting performance in children with autism.
= Autism Diagnostic Interview–Revised;
= Autism Diagnostic Observation Schedule–Generic;
= autism spectrum disorders;
= Diagnostic Interview for Children and Adolescents, 4th edition;
= Diagnostic and Statistical Manual of Mental Disorders, 4th edition;
= full-scale IQ;
= Physical and Neurological Examination for Subtle (Motor) Signs;
= Perceptual Reasoning Indices;
= Wechsler Intelligence Scale for Children–IV.
PMCID: PMC2777071  PMID: 19901244
Brain : a journal of neurology  2007;130(Pt 7):1861-1872.
Human locomotion must be flexible in order to meet varied environmental demands. Alterations to the gait pattern occur on different time scales, ranging from fast, reactive adjustments to slower, more persistent adaptations. A recent study in humans showed the cerebellum to play a key role for slower walking adaptations in inter-limb coordination during split-belt treadmill walking, but not fast reactive changes. It is not known whether cerebral structures are also important in these processes, though some studies of cats have suggested that they are not. In this study, we used a split-belt treadmill walking task to test whether cerebral damage from stroke impairs either type of flexibility. Results showed that stroke involving cerebral structures did not impair either reactive or adaptive abilities and did not disrupt storage of new inter-limb relationships (i.e. after-effects). This suggests that cerebellar interactions with brainstem, rather than cerebral structures, comprise the critical circuit for this type of inter-limb control. Further, the after-effects from a 15-minute adaptation session could temporarily induce symmetry in subjects who demonstrated baseline asymmetry of spatio-temporal gait parameters. In order to re-establish symmetric walking, the choice of which leg is on the fast belt during split-belt walking must be based on the subject’s initial asymmetry. These findings demonstrate that cerebral stroke survivors are indeed able to adapt inter-limb coordination. This raises the possibility that asymmetric walking patterns post-stroke could be remediated utilizing the split-belt treadmill as a long-term rehabilitation strategy.
PMCID: PMC2977955  PMID: 17405765
stroke; motor; locomotion; movement
24.  Split-Belt Treadmill Adaptation Transfers to Overground Walking in Persons Poststroke 
Background and Objective
Following stroke, subjects retain the ability to adapt interlimb symmetry on the split-belt treadmill. Critical to advancing our understanding of locomotor adaptation and its usefulness in rehabilitation is discerning whether adaptive effects observed on a treadmill transfer to walking over ground. We examined whether aftereffects following split-belt treadmill adaptation transfer to overground walking in healthy persons and those poststroke.
Eleven poststroke and 11 age-matched and gender-matched healthy subjects walked over ground before and after walking on a split-belt treadmill. Adaptation and aftereffects in step length and double support time were calculated.
Both groups demonstrated partial transfer of the aftereffects observed on the treadmill (P < .001) to overground walking (P < .05), but the transfer was more robust in the subjects poststroke (P < .05). The subjects with baseline asymmetry after stroke improved in asymmetry of step length and double limb support (P = .06).
The partial transfer of aftereffects to overground walking suggests that some shared neural circuits that control locomotion for different environmental contexts are adapted during split-belt treadmill walking. The larger adaptation transfer from the treadmill to overground walking in the stroke survivors may be due to difficulty adjusting their walking pattern to changing environmental demands. Such difficulties with context switching have been considered detrimental to function poststroke. However, we propose that the persistence of improved symmetry when changing context to overground walking could be used to advantage in poststroke rehabilitation.
PMCID: PMC2811047  PMID: 19307434
Stroke; Locomotion; Rehabilitation; Adaptation
25.  Sensorimotor dysfunction in multiple sclerosis and column-specific magnetization transfer-imaging abnormalities in the spinal cord 
Brain  2009;132(5):1200-1209.
The human spinal cord contains segregated sensory and motor pathways that have been difficult to quantify using conventional magnetic resonance imaging (MRI) techniques. Multiple sclerosis is characterized by both focal and spatially diffuse spinal cord lesions with heterogeneous pathologies that have limited attempts at linking MRI and behaviour. We used a novel magnetization-transfer-weighted imaging approach to quantify damage to spinal white matter columns and tested its association with sensorimotor impairment. We studied 42 participants with multiple sclerosis who each underwent MRI at 3 Tesla and quantitative tests of sensorimotor function. We measured cerebrospinal-fluid-normalized magnetization-transfer signals in the dorsal and lateral columns and grey matter of the cervical cord. We also measured brain lesion volume, cervical spinal cord lesion number and cross-sectional area, vibration sensation, strength, walking velocity and standing balance. We used linear regression to assess the relationship between sensorimotor impairment and MRI abnormalities. We found that the dorsal column cerebrospinal-fluid-normalized magnetization-transfer signal specifically correlated with vibration sensation (R = 0.58, P < 0.001) and the lateral column signal with strength (R = −0.45, P = 0.003). Spinal cord signal measures also correlated with walking and balance dysfunction. A stepwise multiple regression showed that the dorsal column signal and diagnosis subtype alone explained a significant portion of the variance in sensation (R2 = 0.54, P < 0.001), whereas the lateral column signal and diagnosis subtype explained a significant portion of the variance in strength (R2 = 0.30, P < 0.001). These results help to understand the anatomic basis of sensorimotor disability in multiple sclerosis and have implications for testing the effects of neuroprotective and reparative interventions.
PMCID: PMC2677795  PMID: 19297508
strength; sensation; corticospinal tract; dorsal column medial lemniscal tract; magnetic resonance imaging

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