Cerebellar mechanisms for motor learning: Testing predictions from a large-scale computer simulation

The cerebellum is the major brain structure that contributes to our ability to improve movements through learning and experience. We have combined computer simulations with behavioral and lesion studies to investigate how modification of synaptic strength at two different sites within the cerebellum contributes to a simple form of motor learning—Pavlovian conditioning of the eyelid response. These studies are based on the wealth of knowledge about the intrinsic circuitry and physiology of the cerebellum and the straightforward manner in which this circuitry is engaged during eyelid conditioning. Thus, our simulations are constrained by the well-characterized synaptic organization of the cerebellum and further, the activity of cerebellar inputs during simulated eyelid conditioning is based on existing recording data. These simulations have allowed us to make two important predictions regarding the mechanisms underlying cerebellar function, which we have tested and confirmed with behavioral studies. The first prediction describes the mechanisms by which one of the sites of synaptic modification, the granule to Purkinje cell synapses (gr → Pkj) of the cerebellar cortex, could generate two time-dependent properties of eyelid conditioning—response timing and the ISI function. An empirical test of this prediction using small, electrolytic lesions of the cerebellar cortex revealed the pattern of results predicted by the simulations. The second prediction made by the simulations is that modification of synaptic strength at the other site of plasticity, the mossy fiber to deep nuclei synapses (mf → nuc), is under the control of Purkinje cell activity. The analysis predicts that this property should confer mf → nuc synapses with resistance to extinction. Thus, while extinction processes erase plasticity at the first site, residual plasticity at mf → nuc synapses remains. The residual plasticity at the mf → nuc site confers the cerebellum with the capability for rapid relearning long after the learned behavior has been extinguished. We confirmed this prediction using a lesion technique that reversibly disconnected the cerebellar cortex at various stages during extinction and reacquisition of eyelid responses. The results of these studies represent significant progress toward a complete understanding of how the cerebellum contributes to motor learning. ^

Motor learning by field approximation.

We investigated how human subjects adapt to forces perturbing the motion of their ams. We found that this kind of learning is based on the capacity of the central nervous system (CNS) to predict and therefore to cancel externally applied perturbing forces. Our experimental results indicate: (i) that the ability of the CNS to compensate for the perturbing forces is restricted to those spatial locations where the perturbations have been experienced by the moving arm. The subjects also are able to compensate for forces experienced at neighboring workspace locations. However, adaptation decays smoothly and quickly with distance from the locations where disturbances had been sensed by the moving limb. (ii) Our experiments also how that the CNS builds an internal model of the external perturbing forces in intrinsic (muscles and / or joints) coordinates.

The Optimality of Decision Making During Motor Learning

In our daily lives, we often must predict how well we are going to perform in the future based on an evaluation of our current performance and an assessment of how much we will improve with practice. Such predictions can be used to decide whether to invest our time and energy in learning and, if we opt to invest, what rewards we may gain. This thesis investigated whether people are capable of tracking their own learning (i.e. current and future motor ability) and exploiting that information to make decisions related to task reward. In experiment one, participants performed a target aiming task under a visuomotor rotation such that they initially missed the target but gradually improved. After briefly practicing the task, they were asked to select rewards for hits and misses applied to subsequent performance in the task, where selecting a higher reward for hits came at a cost of receiving a lower reward for misses. We found that participants made decisions that were in the direction of optimal and therefore demonstrated knowledge of future task performance. In experiment two, participants learned a novel target aiming task in which they were rewarded for target hits. Every five trials, they could choose a target size which varied inversely with reward value. Although participants’ decisions deviated from optimal, a model suggested that they took into account both past performance, and predicted future performance, when making their decisions. Together, these experiments suggest that people are capable of tracking their own learning and using that information to make sensible decisions related to reward maximization.

The Optimality of Decision Making During Motor Learning

In our daily lives, we often must predict how well we are going to perform in the future based on an evaluation of our current performance and an assessment of how much we will improve with practice. Such predictions can be used to decide whether to invest our time and energy in learning and, if we opt to invest, what rewards we may gain. This thesis investigated whether people are capable of tracking their own learning (i.e. current and future motor ability) and exploiting that information to make decisions related to task reward. In experiment one, participants performed a target aiming task under a visuomotor rotation such that they initially missed the target but gradually improved. After briefly practicing the task, they were asked to select rewards for hits and misses applied to subsequent performance in the task, where selecting a higher reward for hits came at a cost of receiving a lower reward for misses. We found that participants made decisions that were in the direction of optimal and therefore demonstrated knowledge of future task performance. In experiment two, participants learned a novel target aiming task in which they were rewarded for target hits. Every five trials, they could choose a target size which varied inversely with reward value. Although participants’ decisions deviated from optimal, a model suggested that they took into account both past performance, and predicted future performance, when making their decisions. Together, these experiments suggest that people are capable of tracking their own learning and using that information to make sensible decisions related to reward maximization.

Movement Sonification: Effects on Motor Learning beyond Rhythmic Adjustments

Motor learning is based on motor perception and emergent perceptual-motor representations. A lot of behavioral research is related to single perceptual modalities but during last two decades the contribution of multimodal perception on motor behavior was discovered more and more. A growing number of studies indicates an enhanced impact of multimodal stimuli on motor perception, motor control and motor learning in terms of better precision and higher reliability of the related actions. Behavioral research is supported by neurophysiological data, revealing that multisensory integration supports motor control and learning. But the overwhelming part of both research lines is dedicated to basic research. Besides research in the domains of music, dance and motor rehabilitation, there is almost no evidence for enhanced effectiveness of multisensory information on learning of gross motor skills. To reduce this gap, movement sonification is used here in applied research on motor learning in sports. Based on the current knowledge on the multimodal organization of the perceptual system, we generate additional real-time movement information being suitable for integration with perceptual feedback streams of visual and proprioceptive modality. With ongoing training, synchronously processed auditory information should be initially integrated into the emerging internal models, enhancing the efficacy of motor learning. This is achieved by a direct mapping of kinematic and dynamic motion parameters to electronic sounds, resulting in continuous auditory and convergent audiovisual or audio-proprioceptive stimulus arrays. In sharp contrast to other approaches using acoustic information as error-feedback in motor learning settings, we try to generate additional movement information suitable for acceleration and enhancement of adequate sensorimotor representations and processible below the level of consciousness. In the experimental setting, participants were asked to learn a closed motor skill (technique acquisition of indoor rowing). One group was treated with visual information and two groups with audiovisual information (sonification vs. natural sounds). For all three groups learning became evident and remained stable. Participants treated with additional movement sonification showed better performance compared to both other groups. Results indicate that movement sonification enhances motor learning of a complex gross motor skill-even exceeding usually expected acoustic rhythmic effects on motor learning.

The Impact of Motor Learning on Motor Behavior and Cortical Dynamics in a Complex Stressful Social Environment

An economy of effort is a core characteristic of highly skilled motor performance often described as being effortless or automatic. Electroencephalographic (EEG) evaluation of cortical activity in elite performers has consistently revealed a reduction in extraneous associative cortical activity and an enhancement of task-relevant cortical processes. However, this has only been demonstrated under what are essentially practice-like conditions. Recently it has been shown that cerebral cortical activity becomes less efficient when performance occurs in a stressful, complex social environment. This dissertation examines the impact of motor skill training or practice on the EEG cortical dynamics that underlie performance in a stressful, complex social environment. Sixteen ROTC cadets participated in head-to-head pistol shooting competitions before and after completing nine sessions of skill training over three weeks. Spectral power increased in the theta frequency band and decreased in the low alpha frequency band after skill training. EEG Coherence increased in the left frontal region and decreased in the left temporal region after the practice intervention. These suggest a refinement of cerebral cortical dynamics with a reduction of task extraneous processing in the left frontal region and an enhancement of task related processing in the left temporal region consistent with the skill level reached by participants. Partitioning performance into ‘best’ and ‘worst’ based on shot score revealed that deliberate practice appears to optimize cerebral cortical activity of ‘best’ performances which are accompanied by a reduction in task-specific processes reflected by increased high-alpha power, while ‘worst’ performances are characterized by an inappropriate reduction in task-specific processing resulting in a loss of focus reflected by higher high-alpha power after training when compared to ‘best’ performances. Together, these studies demonstrate the power of experience afforded by practice, as a controllable factor, to promote resilience of cerebral cortical efficiency in complex environments.

Implicit and explicit motor learning: Application to children with Autism Spectrum Disorder (ASD)

Aims and objectives: This study aims to determine whether children with Autism Spectrum Disorder (ASD) are capable of learning a motor skill both implicitly and explicitly. Methods: In the present study, 30 boys with ASD, aged 7-11 with IQ average of 81.2, were compared with 32 typical IQ- and age-matched boys on their performance on a serial reaction time task (SRTT). Children were grouped by ASD and typical children and by implicit and explicit learning groups for the SRTT. Results: Implicit motor learning occurred in both children with ASD (p = .02) and typical children (p = .01). There were no significant differences between groups (p = .39). However, explicit motor learning was only observed in typical children (p = .01) not children with ASD (p = .40). There was a significant difference between groups for explicit learning (p = .01). Discussion: The results of our study showed that implicit motor learning is not affected in children with ASD. Implications for implicit and explicit learning are applied to the CO-OP approach of motor learning with children with ASD.

Motor Control and Learning by the State Space Model

A model is presented that deals with problems of motor control, motor learning, and sensorimotor integration. The equations of motion for a limb are parameterized and used in conjunction with a quantized, multi-dimensional memory organized by state variables. Descriptions of desired trajectories are translated into motor commands which will replicate the specified motions. The initial specification of a movement is free of information regarding the mechanics of the effector system. Learning occurs without the use of error correction when practice data are collected and analyzed.

Psychologically Inspired Sensory-Motor Development in Early Robot Learning

Lee, M., Meng, Q. (2005). Psychologically Inspired Sensory-Motor Development in Early Robot Learning. International Journal of Advanced Robotic Systems, 325-334.

Sonification as Concurrent Augmented Feedback for Motor Skill Learning and the Importance of Mapping Design

In recent years, sonification of movement has emerged as a viable method for the provision of feedback in motor learning. Despite some experimental validation of its utility, controlled trials to test the usefulness of sonification in a motor learning context are still rare. As such, there are no accepted conventions for dealing with its implementation. This article addresses the question of how continuous movement information should be best presented as sound to be fed back to the learner. It is proposed that to establish effective approaches to using sonification in this context, consideration must be given to the processes that underlie motor learning, in particular the nature of the perceptual information available to the learner for performing the task at hand. Although sonification has much potential in movement performance enhancement, this potential is largely unrealised as of yet, in part due to the lack of a clear framework for sonification mapping: the relationship between movement and sound. By grounding mapping decisions in a firmer understanding of how perceptual information guides learning, and an embodied cognition stance in general, it is hoped that greater advances in use of sonification to enhance motor learning can be achieved.

Effects of age on learning a spatial motor task in younger and older adults individualizing their knowledge of results schedule

Self-controlled KR practice has revealed that providing participants the opportunity to control their KR is superior for motor learning compared to participants replicating the KR schedule of a self-control participant, without the choice (e.g., yoked). The purpose of the present experiment was two-fold. First, to examine the utility of a self-controlled KR schedule for learning a spatial motor task in younger and older adults and second, to determine whether a self-controlled KR schedule facilitates an increased ability to estimate one’s performance in retention and transfer. Twenty younger adults and 20 older adults practiced in either the self-control or yoked condition and were required to push and release a slide along a confined pathway using their non-dominant hand to a target distance. The retention data revealed that as a function of age, a self-controlled KR schedule facilitated superior retention performance and performance estimations in younger adults compared to their yoked counterparts.

Effects of Proactive and Retroactive Augmented Information on Physiological Responses in Learning a Novel Motor Skill

Previous research has demonstrated superior learning by participants presented with augmented task information retroactively versus proactively (Patterson & Lee, 2008; 2010). Theoretical explanations of these findings are related to the cognitive effort invested by participants during motor skill acquisition. The present study extended previous research by utilizing the physiological index, power spectral analysis of heart rate variability, previously shown to be sensitive to the degree of cognitive effort invested during the performance of a motor task (e.g., increase cognitive effort results in increased LF/HF ratio). Participants were required to learn 18 different key-pressing sequences. As expected, the proactive condition demonstrated superior RS during acquisition, with the retroactive condition demonstrating superior RS during retention. Measures of LF/HF ratio indicated the retroactive participants were investing significantly less cognitive effort in the retention period compared to the proactive participants (p< .05) as a function of learning.