Amy J. Bastian

Cerebellar Contributions to Locomotor Adaptations during Splitbelt Treadmill Walking

Locomotor adaptability ranges from the simple and fast-acting to the complex and long-lasting and is a requirement for successful mobility in an unpredictable environment. Several neural structures, including the spinal cord, brainstem, cerebellum, and motor cortex, have been implicated in the control of various types of locomotor adaptation. However, it is not known which structures control which types of adaptation and the specific mechanisms by which the appropriate adjustments are made. Here, we used a splitbelt treadmill to test cerebellar contributions to two different forms of locomotor adaptation in humans. We found that cerebellar damage does not impair the ability to make reactive feedback-driven motor adaptations, but significantly disrupts predictive feedforward motor adaptations during splitbelt treadmill locomotion. Our results speak to two important aspects of locomotor control. First, we have demonstrated that different levels of locomotor adaptability are clearly dissociable. Second, the cerebellum seems to play an essential role in predictive but not reactive locomotor adjustments. We postulate that reactive adjustments may instead be predominantly controlled by lower neural centers, such as the spinal cord or brainstem.

Learning to predict the future: the cerebellum adapts feedforward movement control.

The role of the cerebellum in motor control and learning has been largely inferred from the effects of cerebellar damage. Recent work shows that cerebellar damage produces greater impairment of movements that require predictive as opposed to reactive control. This dissociation is consistent across many different types of movement. Predictive control is crucial for fast and ballistic movements, but impaired prediction can also affect slow movements, because of increased reliance on time-delayed feedback signals. The new findings are compatible with theories of cerebellar function, but still do not resolve whether the cerebellum operates by predicting the optimal motor commands or future sensory states. Prediction mechanisms must be learned and maintained through comparisons between predicted and observed outcomes. New results show that not all such error information is equivalent in driving cerebellar learning.

Cerebellar Control of Balance and Locomotion

The cerebellum is important for movement control and plays a particularly crucial role in balance and locomotion. As such, one of the most characteristic signs of cerebellar damage is walking ataxia. It is not known how the cerebellum normally contributes to walking, although recent work suggests that it plays a role in the generation of appropriate patterns of limb movements, dynamic regulation of balance, and adaptation of posture and locomotion through practice. The purpose of this review is to examine mechanisms of cerebellar control of balance and locomotion, emphasizing studies of humans and other animals. Implications for rehabilitation are also considered.

Adaptation reveals independent control networks for human walking

Human walking is remarkably adaptable on short and long timescales. We can immediately transition between directions and gait patterns, and we can adaptively learn accurate calibrations for different walking contexts. Here we studied the degree to which different motor patterns can adapt independently. We used a split-belt treadmill to adapt the right and left legs to different speeds and in different directions (forward versus backward). To our surprise, adults could easily walk with their legs moving in opposite directions. Analysis of aftereffects showed that walking adaptations are stored independently for each leg and do not transfer across directions. Thus, there are separate functional networks controlling forward and backward walking in humans, and the circuits controlling the right and left legs can be trained individually. Such training could provide a new therapeutic approach for correcting various walking asymmetries.

Sensory and motor deficits in children with cerebral palsy born preterm correlate with diffusion tensor imaging abnormalities in thalamocortical pathways

Aim  Cerebral palsy (CP) is frequently linked to white matter injury in children born preterm. Diffusion tensor imaging (DTI) is a powerful technique providing precise identification of white matter microstructure. We investigated the relationship between DTI‐observed thalamocortical (posterior thalamic radiation) injury, motor (corticospinal tract) injury, and sensorimotor function.

Size of Error Affects Cerebellar Contributions to Motor Learning

Small errors may affect the process of learning in a fundamentally different way than large errors. For example, adapting reaching movements in response to a small perturbation produces generalization patterns that are different from large perturbations. Are distinct neural mechanisms engaged in response to large versus small errors? Here, we examined the motor learning process in patients with severe degeneration of the cerebellum. Consistent with earlier reports, we found that the patients were profoundly impaired in adapting their motor commands during reaching movements in response to large, sudden perturbations. However, when the same magnitude perturbation was imposed gradually over many trials, the patients showed marked improvements, uncovering a latent ability to learn from errors. On sudden removal of the perturbation, the patients exhibited aftereffects that persisted much longer than did those in healthy controls. That is, despite cerebellar damage, the brain maintained the ability to learn from small errors and the motor memory that resulted from this learning was strongly resistant to change. Of note was the fact that on completion of learning, the motor output of the cerebellar patients remained distinct from healthy controls in terms of its temporal characteristics. Therefore cerebellar degeneration impaired the ability to learn from large-magnitude errors, but had a lesser impact on learning from small errors. The neural basis of motor learning in response to small and large errors appears to be distinct.

Blood flow responses to deep brain stimulation of thalamus

Background and ObjectiveDeep brain stimulation (DBS) of the ventral intermediate nucleus of the thalamus (VIM) provides remarkable relief of tremor in the limbs contralateral to the side of the brain stimulated. The benefits have been sufficiently dramatic that this is now an accepted clinical treatment of essential as well as other forms of tremor. Despite this clinical benefit, the mechanism of action of DBS remains unknown. In this investigation, we sought to determine the effects of VIM DBS on neuronal function. MethodsThe authors used PET measurements of qualitative regional cerebral blood flow in patients with essential tremor to determine the effects of DBS in the left VIM. Each subject had four to six scans with the arms at rest and DBS turned either on or off during alternate scans. Continuous physiologic monitoring revealed no tremor during any of the scans. The PET images from each subject were aligned, averaged, and coregistered to a standard image oriented in stereotactic space. ResultsThe authors used subtraction image analysis with statistical parametric mapping methods and a restricted volume search to identify a significantly increased flow response at the site of stimulation in thalamus. An exploratory analysis revealed increased flow in ipsilateral supplementary motor area, a region that receives afferents from VIM. ConclusionsThe increased blood flow at terminal fields of thalamocortical projections suggests that DBS stimulates and does not inactivate projection neurons in VIM thalamus.

Thinking About Walking: Effects of Conscious Correction Versus Distraction on Locomotor Adaptation

Control of the human walking pattern normally requires little thought, with conscious control used only in the face of a challenging environment or a perturbation. We have previously shown that people can adapt spatial and temporal aspects of walking to a sustained perturbation generated by a split-belt treadmill. Here we tested whether conscious correction of walking, versus distraction from it, modifies adaptation. Conscious correction of stepping may expedite the adaptive process and help to form a new walking pattern. However, because walking is normally an automatic process, it is possible that conscious effort could interfere with adaptation, whereas distraction might improve it by removing competing voluntary control. Three groups of subjects were studied: a control group was given no specific instructions, a conscious correction group was instructed how to step and given intermittent visual feedback of stepping during adaptation, and a distraction group performed a dual-task during adaptation. After adaptation, retention of aftereffects was assessed in all groups during normal treadmill walking without conscious effort, feedback, or distraction. We found that conscious correction speeds adaptation, whereas distraction slows it. Subjects trained with distraction retained aftereffects longest, suggesting that the training used during adaptation predicts the time course of deadaptation. An unexpected finding was that these manipulations affected the adaptation rate of spatial but not temporal elements of walking. Thus conscious processes can preferentially access the spatial walking pattern. It may be that spatial and temporal controls of locomotion are accessible through distinct neural circuits, with the former being most sensitive to conscious effort or distraction.

Reach Adaptation: What Determines Whether We Learn an Internal Model of the Tool or Adapt the Model of Our Arm?

We make errors when learning to use a new tool. However, the cause of error may be ambiguous: is it because we misestimated properties of the tool or of our own arm? We considered a well-studied adaptation task in which people made goal-directed reaching movements while holding the handle of a robotic arm. The robot produced viscous forces that perturbed reach trajectories. As reaching improved with practice, did people recalibrate an internal model of their arm, or did they build an internal model of the novel tool (robot), or both? What factors influenced how the brain solved this credit assignment problem? To investigate these questions, we compared transfer of adaptation between three conditions: catch trials in which robot forces were turned off unannounced, robot-null trials in which subjects were told that forces were turned off, and free-space trials in which subjects still held the handle but watched as it was detached from the robot. Transfer to free space was 40% of that observed in unannounced catch trials. We next hypothesized that transfer to free space might increase if the training field changed gradually, rather than abruptly. Indeed, this method increased transfer to free space from 40 to 60%. Therefore although practice with a novel tool resulted in formation of an internal model of the tool, it also appeared to produce a transient change in the internal model of the subjects arm. Gradual changes in the tools dynamics increased the extent to which the nervous system recalibrated the model of the subjects own arm.

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. Methods. 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. Results. 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). Conclusions. 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.