Musical Sophistication and Multilingualism: Effects on Arcuate Fasciculus Characteristics.

The processing of auditory stimuli which are structured in time is thought to involve the arcuate fasciculus, the white matter tract which connects the temporal cortex and the inferior frontal gyrus. Research has indicated effects of both musical and language experience on the structural characteristics of the arcuate fasciculus. Here, we investigated in a sample of n = 84 young adults whether continuous conceptualizations of musical and multilingual experience related to structural characteristics of the arcuate fasciculus, measured using diffusion tensor imaging. Probabilistic tractography was used to identify the dorsal and ventral parts of the white matter tract. Linear regressions indicated that different aspects of musical sophistication related to the arcuate fasciculus' volume (emotional engagement with music), volumetric asymmetry (musical training and music perceptual abilities), and fractional anisotropy (music perceptual abilities). Our conceptualization of multilingual experience, accounting for participants' proficiency in reading, writing, understanding, and speaking different languages, was not related to the structural characteristics of the arcuate fasciculus. We discuss our results in the context of other research on hemispheric specializations and a dual-stream model of auditory processing.

A kinematically complex multi-articular motor skill for investigating implicit motor learning.

Here we present a task developed to probe implicit learning of a complex motor skill. This task addresses limitations related to task complexity noted in the literature for methods investigating implicit motor learning, namely the serial reaction time task and continuous tracking task. Specifically, the serial reaction time task is limited by the kinematic simplicity of the required movement and the continuous tracing task faces time-on-task confounds and limitations in the control of task difficulty. The task presented herein addresses these issues by employing a kinematically complex multi-articular movement that controls factors that contribute to task difficulty: stimulus animation velocity and trajectory complexity. Accordingly, our objective was to validate the use of this task in probing implicit motor learning, hypothesizing that participants would learn one of the repeating stimuli implicitly. Participants engaged in six blocks of training whereby they first observed and then reproduced a seemingly random complex trajectory. Repeated trajectories were embedded amongst random trajectories. In line with the hypothesis, error for the repeated trajectories was decreased in comparison to that observed for the random trajectories and 73% of participants were unable to identify one of the repeated trajectories, demonstrating the occurrence of implicit learning. While the task requires minor alteration to optimize learning, ultimately the findings underline the task's potential to investigate implicit learning of a complex motor skill.

What we imagine learning from watching others: how motor imagery modulates competency perceptions resulting from the repeated observation of a juggling action.

Although motor learning can occur from observing others perform a motor skill (action observation; AO), observers' confidence in their own ability to perform the skill can be falsely increased compared to their actual ability. This illusion of motor competence (i.e., 'over-confidence') may arise because the learner does not gain access to sensory feedback about their own performance-a source of information that can help individuals understand their veridical motor capabilities. Unlike AO, motor imagery (MI; the mental rehearsal of a motor skill) is thought to be linked to an understanding of movement consequences and kinaesthetic information. MI may thus provide the learner with movement-related diagnostic information, leading to greater accuracy in assessing ability. The present study was designed to evaluate the effects of MI when paired with AO in assessments of one's own motor capabilities in an online observation task. Two groups rated their confidence in performing a juggling task following repeated observations of the action without MI (OBS group; n = 45) or with MI following observation (OBS+MI; n = 39). As predicted, confidence increased with repeated observation for both groups, yet increased to a greater extent in the OBS relative to the OBS+MI group. The addition of MI appeared to reduce confidence that resulted from repeated AO alone. Data support the hypothesis that AO and MI are separable and that MI allows better access to sensory information than AO. However, further research is required to assess changes in confidence that result from MI alone and motor execution.

Learning motor actions via imagery-perceptual or motor learning?

It is well accepted that repeatedly imagining oneself acting without any overt behavior can lead to learning. The prominent theory accounting for why imagery practice is effective, motor simulation theory, posits that imagined action and overt action are functionally equivalent, the exception being activation of the end effector. If, as motor simulation theory states, one can compile the goal, plan, motor program and outcome of an action during imagined action similar to overt action, then learning of novel skills via imagery should proceed in a manner equivalent to that of overt action. While the evidence on motor simulation theory is both plentiful and diverse, it does not explicitly account for differences in neural and behavioural findings between imagined and overt action. In this position paper, we briefly review theoretical accounts to date and present a perceptual-cognitive theory that accounts for often observed outcomes of imagery practice. We suggest that learning by way of imagery reflects perceptual-cognitive scaffolding, and that this 'perceptual' learning transfers into 'motor' learning (or not) depending on various factors. Based on this theory, we characterize consistently reported learning effects that occur with imagery practice, against the background of well-known physical practice effects and show that perceptual-cognitive scaffolding is well-suited to explain what is being learnt during imagery practice.

Examining the role of the supplementary motor area in motor imagery-based skill acquisition.

Motor imagery (MI) and physical practice (PP) have been seen as parallel processes that can drive acquisition of motor skills. Emerging evidence, however, suggests these two processes may be fundamentally different, whereby MI-based motor skill acquisition relies more on effector-independent encoding of movement relative to PP. This alternate view is supported by evidence where real and virtual lesions to brain areas involved in visuospatial processing impair MI-based skill acquisition, and via behavioural studies showing perceptual, but not motor, transfer impairs skill acquisition via MI whereas this effect is reversed in PP. This study further investigated the degree to which MI utilizes effector-independent encoding of movement by investigating the role of the supplementary motor area (SMA), an area involved in perceptual to motor transformations, in MI-based motor skill acquisition. Sixty-four participants completed a serial reaction time paradigm following assignment to one of four groups based on training modality (MI or PP) and stimulation type (sham stimulation or continuous theta burst stimulation to inhibit the SMA). Faster reaction times (RTs) to elements of a repeated sequence in comparison to randomly generated elements indicated that sequence-specific learning occurred. Learning occurred in both PP and MI, with the magnitude of learning significantly smaller in MI. Inhibitory stimulation impaired learning in both modalities. In the context of a framework that distinguishes effector-independent and -dependent components of learning, these findings indicate the SMA plays a role in developing motor chunks in both PP and MI facilitating effector-independent learning in both modalities.

Neural and Behavioral Outcomes Differ Following Equivalent Bouts of Motor Imagery or Physical Practice.

Despite its reported effectiveness for the acquisition of motor skills, we know little about how motor imagery (MI)-based brain activation and performance evolves when MI (the imagined performance of a motor task) is used to learn a complex motor skill compared to physical practice (PP). The current study examined changes in MI-related brain activity and performance driven by an equivalent bout of MI- or PP-based training. Participants engaged in 5 days of either MI or PP of a dart-throwing task. Brain activity (via fMRI) and performance-related outcomes were obtained using a pre/post/retention design. Relative to PP, MI-based training did not drive robust changes in brain activation and was inferior for realizing improvements in performance: Greater activation in regions critical to refining the motor program was observed in the PP versus MI group posttraining, and relative to those driven via PP, MI led only to marginal improvements in performance. Findings indicate that the modality of practice (i.e., MI vs. PP) used to learn a complex motor skill manifests as differences in both resultant patterns of brain activity and performance. Ultimately, by directly comparing brain activity and behavioral outcomes after equivalent training through MI versus PP, this work provides unique knowledge regarding the neural mechanisms underlying learning through MI.

Experience modulates motor imagery-based brain activity.

Whether or not brain activation during motor imagery (MI), the mental rehearsal of movement, is modulated by experience (i.e. skilled performance, achieved through long-term practice) remains unclear. Specifically, MI is generally associated with diffuse activation patterns that closely resemble novice physical performance, which may be attributable to a lack of experience with the task being imagined vs. being a distinguishing feature of MI. We sought to examine how experience modulates brain activity driven via MI, implementing a within- and between-group design to manipulate experience across tasks as well as expertise of the participants. Two groups of 'experts' (basketball/volleyball athletes) and 'novices' (recreational controls) underwent magnetoencephalography (MEG) while performing MI of four multi-articular tasks, selected to ensure that the degree of experience that participants had with each task varied. Source-level analysis was applied to MEG data and linear mixed effects modelling was conducted to examine task-related changes in activity. Within- and between-group comparisons were completed post hoc and difference maps were plotted. Brain activation patterns observed during MI of tasks for which participants had a low degree of experience were more widespread and bilateral (i.e. within-groups), with limited differences observed during MI of tasks for which participants had similar experience (i.e. between-groups). Thus, we show that brain activity during MI is modulated by experience; specifically, that novice performance is associated with the additional recruitment of regions across both hemispheres. Future investigations of the neural correlates of MI should consider prior experience when selecting the task to be performed.

Motor imagery-based skill acquisition disrupted following rTMS of the inferior parietal lobule.

Motor imagery (MI), the mental rehearsal of motor tasks, has promise as a therapy in post-stroke rehabilitation. The potential effectiveness of MI is attributed to the facilitation of plasticity in numerous brain regions akin to those recruited for physical practice. It is suggested, however, that MI relies more heavily on regions commonly affected post-stroke, including left hemisphere parietal regions involved in visuospatial processes. However, the impact of parietal damage on MI-based skill acquisition that underlies rehabilitation remains unclear. Here, we examine the contribution of the left inferior parietal lobule (IPL) to MI using inhibitory transcranial magnetic stimulation (TMS) and an MI-based implicit sequence learning (ISL) paradigm. Participants (N = 27) completed the MI-based ISL paradigm after receiving continuous theta burst stimulation to the left IPL (TMS), or with the coil angled away from the scalp (sham). Reaction time differences (dRT) and effect sizes between implicit and random sequences assessed success of MI-based learning. Mean dRT for the sham group was 36.1 ± 28.2 ms (d = 0.71). Mean dRT in the TMS group was 7.7 ± 38.5 ms (d = 0.11). These results indicate that inhibition of the left IPL impaired MI-based learning. We conclude that the IPL and likely the visuospatial processes it mediates are critical for MI performance and thus MI-based skill acquisition or learning. Ultimately, these findings have implications for the use of MI in post-stroke rehabilitation.

Laterality of brain activity during motor imagery is modulated by the provision of source level neurofeedback.

Motor imagery (MI) may be effective as an adjunct to physical practice for motor skill acquisition. For example, MI is emerging as an effective treatment in stroke neurorehabilitation. As in physical practice, the repetitive activation of neural pathways during MI can drive short- and long-term brain changes that underlie functional recovery. However, the lack of feedback about MI performance may be a factor limiting its effectiveness. The provision of feedback about MI-related brain activity may overcome this limitation by providing the opportunity for individuals to monitor their own performance of this endogenous process. We completed a controlled study to isolate neurofeedback as the factor driving changes in MI-related brain activity across repeated sessions. Eighteen healthy participants took part in 3 sessions comprised of both actual and imagined performance of a button press task. During MI, participants in the neurofeedback group received source level feedback based on activity from the left and right sensorimotor cortex obtained using magnetoencephalography. Participants in the control group received no neurofeedback. MI-related brain activity increased in the sensorimotor cortex contralateral to the imagined movement across sessions in the neurofeedback group, but not in controls. Task performance improved across sessions but did not differ between groups. Our results indicate that the provision of neurofeedback during MI allows healthy individuals to modulate regional brain activity. This finding has the potential to improve the effectiveness of MI as a tool in neurorehabilitation.

Short term effects of contralateral tendon vibration on motor unit discharge rate variability and force steadiness in people with Parkinson's disease.

Background: Vibration of one limb affects motor performance of the contralateral limb, and this may have clinical implications for people with lateralized motor impairments through vibration-induced increase in cortical activation, descending neural drive, or spinal excitability. Objective: The objective of this study was to evaluate the effects of acute biceps brachii tendon vibration on force steadiness and motor unit activity in the contralateral limb of persons with Parkinson's disease. Methods: Ten participants with mild to moderate Parkinson's disease severity performed a ramp, hold and de-ramp isometric elbow flexion at 5% of maximum voluntary contraction with the more-affected arm while vibration was applied to the distal biceps brachii tendon on the contralateral, less-affected arm. Using intramuscular fine wire electrodes, 33 MUs in the biceps brachii were recorded across three conditions (baseline, vibration, and post-vibration). Motor unit recruitment & derecruitment thresholds, discharge rates & variability, and elbow flexion force steadiness were compared between conditions with and without vibration. Results: Coefficient of variation of force and discharge rate variability decreased 37 and 17%, respectively in post-vibration compared with baseline and vibration conditions. Although the motor unit discharge rates did not differ between conditions the total number of motor units active at rest after de-ramp were fewer in the post-vibration condition. Conclusion: Contralateral tendon vibration reduces MU discharge rate variability and enhances force control on the more affected side in persons with Parkinson's disease.

Frontoparietal function and underlying structure reflect capacity for motor skill acquisition during healthy aging.

While capacity for motor skill acquisition changes with healthy aging, there has been little consideration of how age-related changes in brain function or baseline brain structure support motor skill acquisition. We examined: (1) age-dependent changes in functional reorganization related to frontoparietal regions during motor skill acquisition, and (2) whether capacity for motor skill acquisition relates to baseline white matter microstructure in frontoparietal tracts. Healthy older and younger adults engaged in 4 weeks of skilled motor practice. Resting-state functional connectivity (rsFC) assessed functional reorganization before and after practice. Diffusion tensor imaging indexed microstructure of a frontoparietal tract at baseline, generated by rsFC seeds. Motor skill acquisition was associated with decreases in rsFC in healthy older adults and increases in rsFC in healthy younger adults. Frontoparietal tract microstructure was lower in healthy older versus younger adults, yet it was negatively associated with rate of skill acquisition regardless of group. Findings indicate that age-dependent alterations in frontoparietal function and baseline structure of a frontoparietal tract reflect capacity for motor skill acquisition.

Gamified Practice Improves Paretic Arm Motor Behavior in Individuals With Stroke.

BACKGROUND: Stroke is a heterogeneous condition, making choice of treatment, and determination of how to structure rehabilitation outcomes difficult. Individualized goal-directed and repetitive physical practice is an important determinant of motor learning. Yet, many investigations of motor learning after stroke deliver task practice without consideration of individual capability of the learner. OBJECTIVE: We developed a gamified arm rehabilitation task for people with stroke that is personalized to individual capacity for paretic arm movement, provides a high dose of practice, progresses through increasingly difficulty levels that are dependent on the performance of the individual, and is practiced in an engaging environment. The objectives of the current study were to determine if 10 days of gamified, object intercept training using the paretic arm would improve arm movement speed and clinical outcome measures of impairment or function. METHODS: Individuals with chronic stroke and age-matched controls engaged in 10 days of gamified, skilled motor practice of a semi-immersive virtual reality-based intercept and release task. The paretic arm was assessed using the Fugl-Meyer Assessment (motor impairment) and Wolf Motor Function Test (motor function) before and after training. RESULTS: Both groups showed faster arm movement speed with practice; individuals with stroke demonstrated reduced paretic arm motor impairment and increased function after the intervention. Age and sex (for both groups), and time post-stroke were not related to changes in movement speed. CONCLUSIONS: Findings indicate that gamified motor training positively affects paretic arm motor behavior in individuals with mild to severe chronic stroke.

Guidelines for reporting action simulation studies (GRASS): Proposals to improve reporting of research in motor imagery and action observation.

Researchers from multiple disciplines have studied the simulation of actions through motor imagery, action observation, or their combination. Procedures used in these studies vary considerably between research groups, and no standardized approach to reporting experimental protocols has been proposed. This has led to under-reporting of critical details, impairing the assessment, replication, synthesis, and potential clinical translation of effects. We provide an overview of issues related to the reporting of information in action simulation studies, and discuss the benefits of standardized reporting. We propose a series of checklists that identify key details of research protocols to include when reporting action simulation studies. Each checklist comprises A) essential methodological details, B) essential details that are relevant to a specific mode of action simulation, and C) further points that may be useful on a case-by-case basis. We anticipate that the use of these guidelines will improve the understanding, reproduction, and synthesis of studies using action simulation, and enhance the translation of research using motor imagery and action observation to applied and clinical settings.

Implicit Adaptation Processes Promoted by Immediate Offline Visual and Numeric Feedback.

In adaptation learning, visual feedback impacts how adaptation proceeds. With concurrent feedback, a more implicit/feedforward process is thought to be engaged, compared to feedback after movement, which promotes more explicit processes. Due to discrepancies across studies, related to timing and type of visual feedback, we isolated these conditions here. Four groups (N = 52) practiced aiming under rotated feedback conditions; feedback was provided concurrently, immediately after movement (visually or numerically), or visually after a 3 s delay. All groups adapted and only delayed feedback attenuated implicit adaptation as evidenced by post-practice after-effects. Contrary to some suggestions, immediately presented offline and numeric feedback resulted in implicit after-effects, potentially due to comparisons between feedforward information and seen or imagined feedback.

Disrupting somatosensory processing impairs motor execution but not motor imagery.

While motor imagery (MI) is thought to be 'functionally equivalent' with motor execution (ME), the equivalence of feedforward and feedback mechanisms between the two modalities is unexplored. Here, we tested the equivalence of these mechanisms between MI and ME via two experiments designed to probe the role of somatosensory processing (Exp 1), and cognitive processing (Exp 2). All participants were engaged in a previously established force-matching task adapted for MI. A reference force was applied (on scale of 1-10, with higher numbers indicative of greater force) to one index finger while participants matched the force with their opposite index finger via ME or MI (control conditions). Participants then rated the force on the same scale of 1-10. Exp 1: Participants (N = 27) performed the task with tactile stimulation (ME+TAC, MI+TAC) in addition to control conditions. Exp 2: Participants (N = 26) performed the task in dual-task conditions (ME+COG, MI+COG) in addition to control conditions. Results indicate that (Exp 1) tactile stimulation impaired performance in ME but not MI. Dual-task conditions (Exp 2) were not shown to impair performance in either practice modality. Findings suggest that while somatosensory processing is critical for ME, it is not for MI. Overall we indicate a functional equivalence between feedforward/back mechanisms in MI and ME may not exist.

Resting-state brain connectivity correlates of musical sophistication.

Introduction: A growing body of research has investigated how performing arts training, and more specifically, music training, impacts the brain. Recent meta-analytic work has identified multiple brain areas where activity varies as a function of levels of musical expertise gained through music training. However, research has also shown that musical sophistication may be high even without music training. Thus, we aim to extend previous work by investigating whether the functional connectivity of these areas relates to interindividual differences in musical sophistication, and to characterize differences in connectivity attributed to performing arts training. Methods: We analyzed resting-state functional magnetic resonance imaging from n = 74 participants, of whom 37 received performing arts training, that is, including a musical instrument, singing, and/or acting, at university level. We used a validated, continuous measure of musical sophistication to further characterize our sample. Following standard pre-processing, fifteen brain areas were identified a priori based on meta-analytic work and used as seeds in separate seed-to-voxel analyses to examine the effect of musical sophistication across the sample, and between-group analyses to examine the effects of performing arts training. Results: Connectivity of bilateral superior temporal gyrus, bilateral precentral gyrus and cerebellum, and bilateral putamen, left insula, and left thalamus varied with different aspects of musical sophistication. By including these measures of these aspects as covariates in post hoc analyses, we found that connectivity of the right superior temporal gyrus and left precentral gyrus relate to effects of performing arts training beyond effects of individual musical sophistication. Discussion: Our results highlight the potential role of sensory areas in active engagement with music, the potential role of motor areas in emotion processing, and the potential role of connectivity between putamen and lingual gyrus in general musical sophistication.

Modality of practice modulates resting state connectivity during motor learning.

When bookending skilled motor practice, changes in resting state functional magnetic resonance imaging (rs-fMRI; used to characterise synchronized patterns of activity when the brain is at rest) reflect functional reorganization that supports motor memory consolidation and learning. Despite its use in practice in numerous domains, the neural mechanisms underlying motor memory consolidation and learning that result from motor imagery practice (MIP) relative to physical practice are not well understood. The current study examined how rs-fMRI is modulated by skilled motor practice that results through either MIP or physical practice. Two groups of participants engaged in five days of MIP or physical practice of a dart throwing task. Performance and rs-fMRI were captured before and after training. Relative to physical practice, where focal changes in rs-fMRI within a cerebellar-cortical network were observed, MIP stimulated widespread changes in rs-fMRI within a frontoparietal network encompassing bilateral regions. Findings indicate functional reorganization that supports motor memory consolidation and learning is not equivalent across practice modalities. Ultimately, this work provides new information regarding the unique neural underpinnings MIP relies on to drive motor memory consolidation and learning.

In vivo myelin imaging and tissue microstructure in white matter hyperintensities and perilesional white matter.

White matter hyperintensities negatively impact white matter structure and relate to cognitive decline in aging. Diffusion tensor imaging detects changes to white matter microstructure, both within the white matter hyperintensity and extending into surrounding (perilesional) normal-appearing white matter. However, diffusion tensor imaging markers are not specific to tissue components, complicating the interpretation of previous microstructural findings. Myelin water imaging is a novel imaging technique that provides specific markers of myelin content (myelin water fraction) and interstitial fluid (geometric mean T2). Here we combined diffusion tensor imaging and myelin water imaging to examine tissue characteristics in white matter hyperintensities and perilesional white matter in 80 individuals (47 older adults and 33 individuals with chronic stroke). To measure perilesional normal-appearing white matter, white matter hyperintensity masks were dilated in 2 mm segments up to 10 mm in distance from the white matter hyperintensity. Fractional anisotropy, mean diffusivity, myelin water fraction, and geometric mean T2 were extracted from white matter hyperintensities and perilesional white matter. We observed a spatial gradient of higher mean diffusivity and geometric mean T2, and lower fractional anisotropy, in the white matter hyperintensity and perilesional white matter. In the chronic stroke group, myelin water fraction was reduced in the white matter hyperintensity but did not show a spatial gradient in perilesional white matter. Across the entire sample, white matter metrics within the white matter hyperintensity related to whole-brain white matter hyperintensity volume; with increasing white matter hyperintensity volume there was increased mean diffusivity and geometric mean T2, and decreased myelin water fraction in the white matter hyperintensity. Normal-appearing white matter adjacent to white matter hyperintensities exhibits characteristics of a transitional stage between healthy white matter and white matter hyperintensities. This effect was observed in markers sensitive to interstitial fluid, but not in myelin water fraction, the specific marker of myelin concentration. Within the white matter hyperintensity, interstitial fluid was higher and myelin concentration was lower in individuals with more severe cerebrovascular disease. Our data suggests white matter hyperintensities have penumbra-like effects in perilesional white matter that specifically reflect increased interstitial fluid, with no changes to myelin concentration. In contrast, within the white matter hyperintensity there are varying levels of demyelination, which vary based on the severity of cerebrovascular disease. Diffusion tensor imaging and myelin imaging may be useful clinical markers to predict white matter hyperintensity formation, and to stage neuronal damage within white matter hyperintensities.

Probing the Effect of Block Duration on Corticospinal Excitability during Motor Imagery Performance.

Considerable evidence exists related to the behavioral outcomes of motor imagery-based training (MI). Comparatively, there is a relative gap in the literature on how corticospinal excitability, a precursor for experience-dependent plasticity, changes over the course of an MI session, and more specifically if there is an effect of varying the duration of the blocks in which MI is performed. As such, we probed corticospinal excitability during MI, whereby the duration of MI blocks within the session were manipulated yet total exposure to MI was kept constant. Participants performed a total of 24 min of MI of common motor tasks in blocks of 2, 4 or 6 min. Transcranial magnetic stimulation was used to assess corticospinal excitability throughout MI performance. All groups demonstrated increased corticospinal excitability over the session. Owing to a decrease in corticospinal excitability when engaging in 6 min blocks and the variability noted when engaging in 2 min blocks, findings suggest that MI performed in 4 min blocks may be preferable for the generation and maintenance of corticospinal excitability, at least relative to 2 and 6 min blocks. Overall, our findings provide physiological evidence that informs the structure of MI training sessions to optimize their effectiveness.