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How does the time of day of a practice session affect learning of a new motor sequence in the elderly? Participants practiced a given finger tapping sequence either during morning or evening hours. All participants robustly improved performance speed within the session concurrent with a reorganization of the tapping pattern of the sequence. However, evening-trained participants showed additional gains overnight and at 1 wk posttraining; moreover, evening training led to a further reorganization of the tapping pattern offline. A learning experience preceding nocturnal sleep can lead to a task-specific movement routine as an expression of novel “how to” knowledge in the elderly.

White the difficulty of a motor task can act as a stimulus for learning in younger adults, it is unknown how task difficulty interacts with age-related reductions in motor performance and altered brain activation. We examined the effects of task difficulty on motor performance and used electroencephalography (EEG) to probe task-related brain activation after acquisition and 24-h retention of a mirror star-tracing skill in healthy older adults (N = 36, 65-86 years). The results showed that the difficulty of the motor skill affected both the magnitude of motor skill learning and the underlying neural mechanisms. Behavioral data revealed that practicing a motor task at a high difficulty level hindered motor skill consolidation. The EEG data indicated that task difficulty modulated changes in brain activation after practice. Specifically, a decrease in task-related alpha power in frontal and parietal electrodes was only present after practice of the skill at the low and medium, but not the high difficulty level. Taken together, our findings show that a failure to engage neural plasticity through practice of a highdifficulty task is accompanied by reduced motor skill retention in older adults. The data help us better understand how older adults learn new motor skills and might have implications for prescribing motor skill practice according to its difficulty in rehabilitation settings. (c) 2020 The Author(s). Published by Elsevier Ltd on behalf of IBRO. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).

Research has supported two distinct forms of motor skill consolidation that can occur between practice sessions: (1) off-line learning, and (2) memory stabilization. Off-line learning describes performance improvement between practice sessions that is above the gain observed at the end of practice, while memory stabilization describes a gain in performance that is maintained between practice sessions. This study used a Lissajous plot to provide concurrent feedback to train participants to produce a 90° relative phase between the index fingers (flexion/extension motion). Significant improvements in performance emerged after ten trials (5 min) of practice. At the end of training, participants were divided into two delay interval groups before retesting, 2-h and 6-h. The retesting session started with participants performing an interference task (10 trials, 5 min) that required training on a 45° relative phase between the fingers with concurrent feedback from the Lissajous plot. When training with the interference task was completed participants were retested with the 90° relative phase without the Lissajous plot feedback. In the retest of the 90° pattern, a performance loss was found in the 2-h delay group, whereas the 6-h delay group maintained the end of practice performance level. Maintenance of the same level of performance without the Lissajous plot represents memory stabilization of the initially trained 90° pattern. The findings are discussed within the context of current positions regarding procedural consolidation and the coordination dynamics framework wherein action and perception are linked through the informational nature of relative phase.

Time-of-day influences both physical and mental performance. Its impact on motor learning is, however, not well established yet. Here, using a finger tapping-task, we investigated the time-of-day effect on skill acquisition (i.e., immediately after a physical or mental practice session) and consolidation (i.e., 24 hours later). Two groups (one physical and one mental) were trained in the morning (10 a.m.) and two others (one physical and one mental) in the afternoon (3 p.m.). We found an enhancement of motor skill following both types of practice, whatever the time of the day, with a better acquisition for the physical than the mental group. Interestingly, there was a better consolidation for both groups when the training session was scheduled in the afternoon. Overall, our results indicate that the time-of-day positively influences motor skill consolidation and thus must be considered to optimize training protocols in sport and clinical domains to potentiate motor (re)learning.

Transcranial alternating current stimulation (tACS) is the application of subthreshold, sinusoidal current to modulate ongoing brain rhythms related to sensory, motor and cognitive processes. Electrophysiological studies suggested that the effect of tACS applied at an alpha frequency (8–12 Hz) was state-dependent. The effects of tACS, that is, an increase in parieto-occipital electroencephalography (EEG) alpha power and magnetoencephalography (MEG) phase coherence, was only observed when the eyes were open (low alpha power) and not when the eyes were closed (high alpha power). This state-dependency of the effects of alpha tACS might extend to the aging brain characterized by general slowing and decrease in spectral power of the alpha rhythm. We additionally hypothesized that tACS will influence the motor cortex, which is involved in motor skill learning and consolidation. A group of young and old healthy adults performed a serial reaction time task (SRTT) with their right hand before and after the tACS stimulation. Each participant underwent three sessions of stimulation: sham, stimulation applied at the individual participant’s alpha peak frequency or individual alpha peak frequency (iAPF; α-tACS) and stimulation with iAPF plus 2 Hz (α2-tACS) to the left motor cortex for 10 min (1.5 mA). We measured the effect of stimulation on general motor skill (GMS) and sequence-specific skill (SS) consolidation. We found that α-tACS and α2-tACS improved GMS and SS consolidation in the old group. In contrast, α-tACS minimally improved GMS consolidation but impaired SS consolidation in the young group. On the other hand, α2-tACS was detrimental to the consolidation of both skills in the young group. Our results suggest that individuals with aberrant alpha rhythm such as the elderly could benefit more from tACS stimulation, whereas for young healthy individuals with intact alpha rhythm the stimulation could be detrimental.

Huntington’s disease (HD) is a neurodegenerative disease notably characterized by progressive motor symptoms. Although the loss of medium spiny neurons (MSNs) in the striatum has been associated with motor deficits, premanifest patients already present cognitive deficiencies and show early signs of motor disabilities. Here, in a YAC128 HD mouse model, we identified impairment in motor skill consolidation at the age of 11–14 weeks., Huntington’s disease (HD) is a neurodegenerative disease notably characterized by progressive motor symptoms. Although the loss of medium spiny neurons (MSNs) in the striatum has been associated with motor deficits, premanifest patients already present cognitive deficiencies and show early signs of motor disabilities. Here, in a YAC128 HD mouse model, we identified impairment in motor skill consolidation at the age of 11–14 weeks. Using optogenetic stimulation, we found that excitatory synaptic transmission from motor cortex to MSNs located in the dorsolateral part of the striatum (DLS) is altered. Using single pellet reaching task, we observed that while motor skill consolidation is accompanied by a dynamic change in AMPA/NMDA ratio in wild-type (WT) mice, this form of synaptic plasticity does not occur in YAC128 mice. This study not only proposes new meaningful insight in the synaptopathic mechanisms of HD, but also highlights that deficit in motor skill consolidation-dependent synaptic plasticity at motor cortex to DLS synapses represents an early biomarker for HD.

Post-training sleep benefits both declarative and procedural memory consolidation. However, recent research suggests that eyes-closed waking rest may provide a similar benefit. Brokaw et al. (2016), for example, recently demonstrated that verbal declarative memory improved more following a 15 min period of waking rest, in comparison to 15 min of active wake. Here, we used the same procedures to test whether procedural memory similarly benefits from waking rest. Participants were trained on the Motor Sequence Task (MST), followed by a 15 min retention interval during which they either rested with their eyes closed or completed a distractor task. Rest significantly enhanced MST performance, mirroring the effect observed in Brokaw et al. (2016) and demonstrating that waking rest benefits the early stages of procedural memory. An additional group of participants tested 4 h later displayed no effect of rest. Overall, these results suggest that the early MST performance "boost" described in prior studies may depend on post-learning state.

Feedback on motor performance activates the striatum and boosting ventral striatum activation with rewarding feedback during motor training supports the consolidation of the learned skill. Aging is associated with changes of the reward system, including striatal and extrastriatal loss of dopamine receptors. How these changes interact with the blood oxygenation level dependent (BOLD) response is, however, not yet fully understood. While it is known that reward prediction and reward-based decision-making differ between young and elderly healthy adults, the influence of age on the processing of rewarding feedback on motor performance have not been investigated so far. Nineteen young (26.42±2.84years) and 18 elderly (65.39±6.40years) healthy adults performed an arc-tracking task including performance feedback linked to a monetary reward after half of the trials, while undergoing functional magnetic resonance imaging (fMRI). The BOLD effect was compared in three predefined regions of interest: Ventral and dorsal striatum plus primary motor cortex. Our study demonstrates differences in the processing of motor performance related reward between young and elderly healthy adults. While both groups earned similar amounts of money linked to their own performance, the ventral striatal response to the rewarding feedback was higher in the older group. Deficient prediction about the rewarding feedback, a higher motivational status or compensation for a reduced number of dopamine receptors in the elderly might be possible explanations. How this interacts with the reward-induced improvement of motor skill consolidation, as observed in young subjects, has to be clarified.

Gamma-aminobutyric acid (GABA) inhibition shapes motor cortex output, gates synaptic plasticity in the form of long-term potentiation, and plays an important role in motor learning. Remarkably, recent studies have shown that acute cardiovascular exercise can improve motor memory, but the cortical mechanisms are not completely understood. We investigated whether an acute bout of lower-limb high-intensity interval (HIT) exercise could promote motor memory formation in humans through changes in cortical inhibition within the hand region of the primary motor cortex. We used TMS to assess the input–output relationship, along with inhibition involving GABAA and GABAB receptors. Measures were obtained before and after a 20-min session of HIT cycling (exercise group) or rest (control group). We then had the same participants learn a new visuomotor skill and perform a retention test 5 hr later in the absence of sleep. No differences were found in corticomotor excitability or GABAB inhibition; however, synaptic GABAA inhibition was significantly reduced for the exercise group but not the control group. HIT exercise was found to enhance motor skill consolidation. These findings link modification of GABA to improved motor memory consolidation after HIT exercise and suggest that the beneficial effects of exercise on consolidation might not be dependent on sleep.

Region- and pathway-specific plasticity within striatal circuits is critically involved in the acquisition and long-term retention of a new motor skill as it becomes automatized. However, the molecular substrates contributing to this plasticity remain unclear. Here, we examined the expression of the activity-regulated cytoskeleton-associated protein (Arc) in the associative or dorsomedial striatum (DMS) and the sensorimotor or dorsolateral striatum (DLS), as well as in striatonigral and striatopallidal neurons, during different skill learning phases in the accelerating rotarod task. We found that Arc was mainly expressed in the DMS during early motor learning and progressively increased in the DLS during gradual motor skill consolidation. Moreover, Arc was preferentially expressed in striatopallidal neurons early in training and gradually increased in striatonigral neurons later in training. These data demonstrate that in the dorsal striatum, the expression of Arc exhibits a region- and cell-specific transfer during the learning of a motor skill, suggesting a link between striatal Arc expression and motor skill learning in mice.

Here our goal was to determine the magnitude of sleep-related motor skill enhancement. Performance on the finger tapping task (FTT) was evaluated after a 90min daytime nap (n=15) or after quiet wakefulness (n=15). By introducing a slight modification in the formula used to calculate the offline gains we were able to refine the estimated magnitude of sleep׳s effect on motor skills. The raw value of improvement after a nap decreased after this correction (from ~15% to ~5%), but remained significantly higher than the control. These results suggest that sleep does indeed play a role in motor skill consolidation.

Sleep benefits motor memory consolidation. This mnemonic process is thought to be mediated by thalamo-cortical spindle activity during NREM-stage2 sleep episodes as well as changes in striatal and hippocampal activity. However, direct experimental evidence supporting the contribution of such sleep-dependent physiological mechanisms to motor memory consolidation in humans is lacking. In the present study, we combined EEG and fMRI sleep recordings following practice of a motor sequence learning (MSL) task to determine whether spindle oscillations support sleep-dependent motor memory consolidation by transiently synchronizing and coordinating specialized cortical and subcortical networks. To that end, we conducted EEG source reconstruction on spindle epochs in both cortical and subcortical regions using novel deep-source localization techniques. Coherence-based metrics were adopted to estimate functional connectivity between cortical and subcortical structures over specific frequency bands. Our findings not only confirm the critical and functional role of NREM-stage2 sleep spindles in motor skill consolidation, but provide first-time evidence that spindle oscillations [11–17 Hz] may be involved in sleep-dependent motor memory consolidation by locally reactivating and functionally binding specific task-relevant cortical and subcortical regions within networks including the hippocampus, putamen, thalamus and motor-related cortical regions.

A single bout of high-intensity exercise can augment off-line gains in skills acquired during motor practice. It is currently unknown if the type of physical exercise influences the effect on motor skill consolidation. This study investigated the effect of three types of high-intensity exercise following visuomotor skill acquisition on the retention of motor memory in 40 young (25.3 ±3.6 years), able-bodied male participants randomly assigned to one of four groups either performing strength training (STR), circuit training (CT), indoor hockey (HOC) or rest (CON). Retention tests of the motor skill were performed 1 (R1h) and 24 h (R1d) post acquisition. For all exercise groups, mean motor performance scores decreased at R1h compared to post acquisition (POST) level; STR (P = 0.018), CT (P = 0.02), HOC (P = 0.014) and performance scores decreased for CT compared to CON (P = 0.049). Mean performance scores increased from POST to R1d for all exercise groups; STR (P = 0.010), CT (P = 0.020), HOC (P = 0.007) while performance scores for CON decreased (P = 0.043). Changes in motor performance were thus greater for STR (P = 0.006), CT (P

Recent reports have revealed that motor skill learning is impaired if two skills are practiced one after the other, that is before the first skill has had the time to become consolidated. This suggests that motor skills should be practiced in isolation from one another to minimize interference. At the moment, little is known about the effect of practice schedules high in contextual interference on motor skill consolidation. In Experiment 1, we investigated whether a serial practice schedule impairs motor skill consolidation. Participants had to learn two distinct sequences of finger movements (A and B) under either a blocked practice schedule or a serial practice schedule before being retested the following day. A control group also practiced Sequence A only. Our results revealed that a blocked practice schedule led to no interference between the sequences, whereas a serial practice schedule impaired the consolidation of Sequence B. In Experiment 2, we investigated the origin of the interference caused by a serial practice schedule by replacing the physical practice of Sequence A with either the observation of a model performing Sequence A or by asking participants to produce random finger movements. Our results revealed that both tasks interfered with the consolidation of Sequence B. Thus, we suggest that a serial practice schedule impairs motor skill consolidation through a conflict in the brain networks involved in the acquisition of the cognitive representation of the sequence and its execution.

The purpose of this study was to determine whether exposure to specific auditory sequences leads to the induction of new motor memories and to investigate the role of the dorsal premotor cortex (dPMC) in this crossmodal learning process. Fifty-two young healthy non-musicians were familiarized with the sound to key-press mapping on a computer keyboard and tested on their baseline motor performance. Each participant received subsequently either continuous theta burst stimulation (cTBS) or sham stimulation over the dPMC and was then asked to remember a 12-note melody without moving. For half of the participants, the contour of the melody memorized was congruent to a subsequently performed, but never practiced, finger movement sequence (Congruent group). For the other half, the melody memorized was incongruent to the subsequent finger movement sequence (Incongruent group). Hearing a congruent melody led to significantly faster performance of a motor sequence immediately thereafter compared to hearing an incongruent melody. In addition, cTBS speeded up motor performance in both groups, possibly by relieving motor consolidation from interference by the declarative melody memorization task. Our findings substantiate recent evidence that exposure to a movement-related tone sequence can induce specific, crossmodal encoding of a movement sequence representation. They further suggest that cTBS over the dPMC may enhance early offline procedural motor skill consolidation in cognitive states where motor consolidation would normally be disturbed by concurrent declarative memory processes. These findings may contribute to a better understanding of auditory-motor system interactions and have implications for the development of new motor rehabilitation approaches using sound and non-invasive brain stimulation as neuromodulatory tools.

The smooth, coordinated fine motor movements required to play a musical instrument are not only highly valued in our society; they also predict academic success in areas that generalize beyond the motor domain, including reading and math readiness, and verbal abilities. Interestingly, motor skills that overlap with those required to play a musical instrument (e.g., sequential finger tapping) markedly improve (get faster) over a night of sleep, but not after a day spent awake. Here we studied whether individuals who play musical instruments that require fine finger motor skill are better able to learn and consolidate a simple motor skill task compared to those who do not play an instrument, and whether sleep-specific motor skill benefits interact with those imparted by musical experience. We used the motor sequence task (MST), which taps into a core skill learned and used by musicians, namely, the repetition of learned sequences of key presses. Not surprisingly, we found that musicians were faster than non-musicians throughout the learning session, typing more correct sequences per 30-sec trial. In the 12hrs that followed learning we found that sleep and musical experience both led to greater improvement in performance. Surprisingly, musicians retested after a day of wake performed slightly better than non-musicians who had slept between training and retest, suggesting that musicians have the capacity to consolidate a motor skill across waking hours, while non-musicians appear to lack this capacity. These findings suggest that the musically trained brain is optimized for motor skill consolidation across both wake and sleep, and that sleep may simply promote a more effective use of this machinery. In sum, there may be something special about musicians, perhaps a neurophysiological advantage, that leads to both the expected—greater motor speed at learning—and the surprising—greater motor skill improvement over time.

Knowledge of performance can activate the striatum, a key region of the reward system and highly relevant for motivated behavior. Using functional magnetic resonance imaging, striatal activity linked to knowledge of performance was measured during the training of a repetitive arc-tracking task. Knowledge of performance was given after a random selection of trials or after good performance. The third group received knowledge of performance after good performance plus a monetary reward. Skill learning was measured from pre- to post- (acquisition) and from post- to 24h posttraining (consolidation). Our results demonstrate an influence of feedback on motor skill learning. Adding a monetary reward after good performance leads to better consolidation and higher ventral striatal activation than knowledge of performance alone. In turn, rewarding strategies that increase ventral striatal response during training of a motor skill may be utilized to improve skill consolidation.

Purpose Sleep following training can enhance motor skill memory consolidation while chronic sleep disruption can have the converse effect. The aim of this investigation was to explore the relationship between sleep measured by wrist actigraphy, motor skill consolidation and primary motor cortex excitability in young, healthy individuals. Methods Training was a visuospatial finger-tracking task. Dependent measures included tracking skill performance, cortical excitability, measures of sleep, and level of arousal. Assessments occurred pre-training, post-training and at 12 h and 24 h retention. An activity monitor was worn on the wrist during the nights preceding and following training. Results Results indicate that sleep during the night following training was predictive of 1) offline skill consolidation following training (R² = 0.34) and 2) cortical excitability at 24 h follow-up (R² = 0.35) with less time spent awake associated with better skill performance and lower cortical excitability at 24 h follow-up. No difference in measures of sleep was observed between nights of sleep (p > 0.05). Sleep the night before training did not influence skill performance, skill acquisition during training, nor measures of cortical excitability at pre-training assessment. Conclusions These findings suggest a relationship between motor skill development, cortical excitability and sleep following training. These results invite further investigation into the utility of actigraphy as a low-cost, easy-to-administer alternative to polysomnography for short and long-term evaluation of the relationship between sleep, cortical excitability and motor skill learning in healthy and patient populations.

In humans the consolidation of recently learned motor skills is a multi-step process. We previously showed that performance on the finger-tapping task (FTT; i.e. a sequential motor skill) temporarily improves early on, 5-30 min after practice has ended, but not 4 h later. In the absence of any further practice to the task, this early boost in performance was predictive of the performance levels eventually achieved 48 h later, suggesting its functional relevance for long-term memory consolidation [Hotermans, Peigneux, Maertens de Noordhout, Moonen, and Maquet (2006) Early boost and slow consolidation in motor skill learning. Learn. Mem., 13, 580-583]. Here, we focused on the role of the primary motor cortex (M1) in consolidation using repetitive transcranial magnetic stimulation (rTMS) applied immediately before testing at 30 min, 4 or 24 h after practice of the FTT. Immediately after learning, rTMS over M1 depressed the early boost in performance, but did not affect the delayed improvement observed 48 h later. Four and 24 h after practice, rTMS did not disrupt performance anymore. These results suggest that M1 supports performance during the early post-training phase of motor skill consolidation, but is no longer mandatory in the subsequent, delayed stages of consolidation.

Sleep is beneficial for performance across a range of memory tasks in young adults, but whether memories are similarly consolidated in older adults is less clear. Performance benefits have been observed following sleep in older adults for declarative learning tasks, but this benefit may be reduced for non-declarative, motor skill learning tasks. To date, studies of sleep-dependent consolidation of motor learning in older adults are limited to motor sequence tasks. To examine whether reduced sleep-dependent consolidation in older adults is generalizable to other forms of motor skill learning, we examined performance changes over intervals of sleep and wake in young (n = 62) and older adults (n = 61) using a mirror-tracing task, which assesses visuo-motor adaptation learning. Participants learned the task either in the morning or in evening, and performance was assessed following a 12-h interval containing overnight sleep or daytime wake. Contrary to our prediction, both young adults and older adults exhibited sleep-dependent gains in visuo-motor adaptation. There was a correlation between performance improvement over sleep and percent of the night in non-REM stage 2 sleep. These results indicate that motor skill consolidation remains intact with increasing age although this relationship may be limited to specific forms of motor skill learning.

While intervening sleep promotes the consolidation of memory, it is well established that cognitive interference from competing stimuli can impede memory retention. The current study examined changes in motor skill learning across periods of wakefulness with and without competing stimuli, and periods of sleep with and without disruption from external stimuli. A napping study design was adopted where participants (N=44) either had (1) a 30min nap composed of Non-Rapid Eye Movement (NREM) sleep, (2) 30min NREM nap fragmented by audio tone induced arousals, (3) 45min of quiet wakefulness, or (4) 45min of active wakefulness. Measures of subjective sleepiness (KSS), alertness (PVT) and motor skill learning (Sequential Finger Tapping Task, SFTT) were completed in the morning and evening to assess performance pre- and post-nap or wakefulness. Following a practice session, change in motor skill performance was measured over a 10min post training rest interval, as well as following a 7h morning to evening interval comprising one of the four study conditions. A significant offline enhancement in motor task performance (13-23%) was observed following 10min of rest in all conditions. Following the long delay with the intervening nap/wake condition, there were no further offline gains or losses in performance in any sleep (uninterrupted/fragmented) or wake (quiet/active) condition. The current findings suggest that after controlling for offline gains in performance that occur after a brief rest and likely to due to the dissipation of fatigue, the subsequent effect of an intervening sleep or wake period on motor skill consolidation is not significant. Consistent with this null result, the impact of disrupting the sleep episode or manipulating activity during intervening wake also appears to be negligible.

Improvement in motor skill performance is known to continue for at least 24 hr following training, yet the relative contributions of time spent awake and asleep are unknown. Here we provide evidence that a night of sleep results in a 20% increase in motor speed without loss of accuracy, while an equivalent period of time during wake provides no significant benefit. Furthermore, a significant correlation exists between the improved performance overnight and the amount of stage 2 NREM sleep, particularly late in the night. This finding of sleep-dependent motor skill improvement may have important implications for the efficient learning of all skilled actions in humans.

Article About the Authors Metrics Comments Related Content Abstract Introduction Materials and Methods Results Discussion Conclusions Supporting Information Acknowledgments Author Contributions References Reader Comments (0) FiguresAbstractGilles de la Tourette syndrome is a neuropsychiatric disorder characterized by motor and phonic tics that can be considered motor responses to preceding inner urges. It has been shown that Tourette patients have inferior performance in some motor learning tasks and reduced synaptic plasticity induced by transcranial magnetic stimulation. However, it has not been investigated whether altered synaptic plasticity is directly linked to impaired motor skill acquisition in Tourette patients. In this study, cortical plasticity was assessed by measuring motor-evoked potentials before and after paired associative stimulation in 14 Tourette patients (13 male; age 18–39) and 15 healthy controls (12 male; age 18–33). Tic and urge severity were assessed using the Yale Global Tic Severity Scale and the Premonitory Urges for Tics Scale. Motor learning was assessed 45 minutes after inducing synaptic plasticity and 9 months later, using the rotary pursuit task. On average, long-term potentiation-like effects in response to the paired associative stimulation were present in healthy controls but not in patients. In Tourette patients, long-term potentiation-like effects were associated with more and long-term depression-like effects with less severe urges and tics. While motor learning did not differ between patients and healthy controls 45 minutes after inducing synaptic plasticity, the learning curve of the healthy controls started at a significantly higher level than the Tourette patients' 9 months later. Induced synaptic plasticity correlated positively with motor skills in healthy controls 9 months later. The present study confirms previously found long-term improvement in motor performance after paired associative stimulation in healthy controls but not in Tourette patients. Tourette patients did not show long-term potentiation in response to PAS and also showed reduced levels of motor skill consolidation after 9 months compared to healthy controls. Moreover, synaptic plasticity appears to be related to symptom severity.

Preface - NICOLA HODGES and MARK WILLIAMS I. PRESENTING INFORMATION 1. The roles and uses of augmented feedback in motor skill acquisition - RICHARD MAGILL and DAVID ANDERSON 2. Mixing it up a little: How to schedule observational practice - NICOLE ONG and NICOLA HODGES 3. Attentional focus affects movement efficiency - KEITH LOHSE, GABRIELE WULF and REBECCA LEWTHWAITE 4. Advances in implicit motor learning - RICHARD S. W. MASTERS and JAMIE POOLTON II: OPTIMIZING PRACTICE CONDITIONS 5. Scheduling practice - TIMOTHY LEE 6. Mental imagery, action observation and skill learning - AIDAN MORAN, MARK CAMPBELL, PAUL HOLMES and TADHG MACINTYRE 7. Ecological dynamics and motor learning design in sport - KEITH DAVIDS, DUARTE ARAUJO, ROBERT HRISTOVSKI, PEDRO PASSOS AND JIA YI CHOW 8. The representation, production, and transfer of simple and complex movement sequences - CHARLES SHEA and DAVID WRIGHT 9. Physical guidance research: Assisting principles and supporting evidence - NICOLA HODGES and PAUL CAMPAGNARO III: ISSUES IN MOTOR LEARNING 10. Motor learning through a motivational lens - REBECCA LEWTHWAITE and GABRIELE WULF 11. Motor skill consolidation - MAXIME TREMPE and LUC PROTEAU 12. Critical periods, readiness and skill learning - DAVID ANDERSON, RICHARD MAGILL and REGIS THOUVARECQ 13. Mechanisms of skilled joint action performance - TERRY ESKENAZI, ROBRECHT VAN DER WEL and NATALIE SEBANZ 14. Motor skill learning and its neurophysiology - KATIE WADDEN, MICHAEL BORICH and LARA BOYD IV: SKILLED PERFORMANCE 15. The development of skill in sport - JEAN COTE, JENNIFER MURPHY MILLS and BRUCE ABERNETHY 16. Experts have all the time in the world - BRUCE ABERNETHY, DAMIAN FARROW, ADAM GORMAN and DAVID MANN 17. Perceptual expertise: what can be trained? JOE CAUSER, CHRISTOPHER JANELLE, JOAN VICKERS and A. MARK WILLIAMS 18. Embodied cognition: from the playing field to the classroom - CARLY KONTRA, NEIL ALBERT and SIAN BEILOCK 19. Especial skills: generality and specificity in motor learning - GAVIN BRESLIN, RICHARD SCHMIDT and TIMOTHY LEE V: RESEARCH, THEORY & PRACTICE: CHALLENGES & SOLUTIONS 20. Translating Theory into Practice: Working at the 'Coal Face' in the UK! - A. MARK WILLIAMS, PAUL FORD, JOE CAUSER, OLIVER LOGAN and STAFFORD MURRAY 21. Working in the field: A Southern hemisphere perspective - CHRIS BUTTON and DAMIAN FARROW

Sleep is known to contribute to motor memory consolidation. Recent studies have provided evidence that a night of sleep plays a similar functional role following motor imagery (MI), while the simple passage of time does not result in performance gains. Here, we examined the benefits of a daytime nap on motor memory consolidation after MI practice. Participants were trained by MI on an explicitly known sequence of finger movements at 11:00. Half of the participants were then subjected (at 14:00) to either a short nap (10 min of stage 2 sleep) or a long nap (60-90 min, including slow wave sleep and rapid eye movement sleep). We also collected data from both quiet and active rest control groups. All participants remained in the lab until being retested at 16:00. The data revealed that a daytime nap after imagery practice improved motor performance and, therefore, facilitated motor memory consolidation, as compared with spending a similar time interval in the wake state. Interestingly, the results revealed that both short and long naps resulted in similar delayed performance gains. The data might also suggest that the presence of slow wave and rapid eye movement sleep does not provide additional benefits for the sleep-dependent motor skill consolidation following MI practice.

The influence of sleep on motor skill consolidation has been a research topic of increasing interest. In this study, we distinguished general skill learning from sequence-specific learning in a probabilistic implicit sequence learning task (alternating serial reaction time) in young and old adults before and after a 12-h offline interval which did or did not contain sleep (p.m.-a.m. and a.m.-p.m. groups, respectively). The results showed that general skill learning, as assessed via overall reaction time, improved offline in both the young and older groups, with the young group improving more than the old. However, the improvement was not sleep-dependent, in that there was no difference between the a.m.-p.m. and p.m.-a.m. groups. We did not find sequence-specific offline improvement in either age group for the a.m.-either p.m. or p.m.-a.m. groups, suggesting that consolidation of this kind of implicit motor sequence learning may not be influenced by sleep.

Rapid eye movement (REM) sleep has been considered important for the consolidation of memories, particularly of procedural skills. REM sleep, in contrast to slow-wave sleep (SWS), is hallmarked by the high, wake-like activity of the neurotransmitter acetylcholine (ACh), which promotes certain synaptic plastic processes underlying the formation of memories. Here, we show in healthy young men that off-line consolidation of a motor skill during a period of late sleep with high amounts of REM sleep depends essentially on high cholinergic activity. After a 3-h sleep period during the early night to satisfy the need for SWS, subjects learned a procedural finger sequence tapping task and a declarative word-pair learning task. After learning, they received either placebo or a combination of the muscarinic receptor antagonist scopolamine (4 microg/kg bodyweight, intravenously) and the nicotinic receptor antagonist mecamylamine (5 mg, orally), and then slept for another 3 h, ie, the late nocturnal sleep period, which is dominated by REM sleep. Retrieval was tested the following evening. Combined cholinergic receptor blockade significantly impaired motor skill consolidation, whereas word-pair memory remained unaffected. Additional data show that the impairing effect of cholinergic receptor blockade is specific to sleep-dependent consolidation of motor skill and does not occur during a wake-retention interval. Taken together, these results identify high cholinergic activity during late, REM sleep-rich sleep as an essential factor promoting sleep-dependent consolidation of motor skills.

We investigated the acquisition and long-term retention of new skills in patients with cortical (Alzheimer's disease, AD) and subcortical (progressive supranuclear palsy, PSP; Parkinson's disease, PD) degeneration. The motor skill task performance of the PD and PSP patients improved with training, but the improvement disappeared within a few months, whereas AD patients retained learned skills for 3-18 months. The results of our experiments show that subcortical dysfunction induces a retention deficit for newly learned motor skills. Our present study suggests that a normal striatum is necessary for the formation of long-lasting motor skills, and that the striatum plays an important role as a motor skill consolidation system.

A wealth of recent studies support a function of sleep on memory and cognitive processing. At a physiological level, sleep supports memory in a number of ways including neural replay and enhanced plasticity in the context of reduced ongoing input. This paper presents behavioral evidence for sleep’s role in selective remembering and forgetting of declarative memories, in generalization of these memories, and in motor skill consolidation. Recent physiological data reviewed suggests how these behavioral changes might be supported by sleep. Importantly, in reviewing these findings, an integrated view of how distinct sleep stages uniquely contribute to memory processing emerges. This model will be useful in developing future behavioral and physiological studies to test predictions that emerge.

Recently, it has been suggested that the primary motor cortex (M1) plays a critical role in implementing the fast and transient post-training phase of motor skill consolidation, known to yield an early boost in performance. Whether a comparable early boost in performance occurs following motor imagery (MIM) training is still unknown. To address this issue, two groups of subjects learned a finger tapping sequence either by MIM or physical practice (PP). In both groups, performance increased significantly in the post-training phase when compared with the pre-training phase and further increased after a 30 min resting period, indicating that both MIM and PP trainings were equally efficient and induced an early boost in motor performance. This conclusion was corroborated by the results of an additional control group. In a second experiment, we then investigated the causal role of M1 in implementing the early boost process resulting from MIM training. To do so, we inhibited M1 by applying a continuous theta-burst stimulation (cTBS) in healthy volunteers just after they learnt, by MIM, the same finger-tapping task as in Experiment #1. As a control, cTBS was applied over the vertex of subjects who underwent the same experiment. We found that cTBS applied over M1 selectively abolished the early boost process subsequent to MIM training. Altogether, the present study provides evidence that MIM practice induces an early boost in performance and demonstrates that M1 is causally involved in this process. These findings further divulge some degree of behavioral and neuronal similitude between MIM and PP.

The retention of a new sequential motor skill relies on repeated practice and subsequent consolidation in the absence of active skill practice. While the early phase of skill acquisition remains relatively unaffected in older adults, posttraining consolidation appears to be selectively impaired by advancing age. Motor learning is associated with posttraining changes of oscillatory alpha and beta neuronal activities in the motor cortex. However, whether or not these oscillatory dynamics relate to posttraining consolidation and how they relate to the age-specific impairment of motor consolidation in older adults remains elusive. Transcranial alternating current stimulation (tACS) is a noninvasive brain stimulation technique capable of modulating such neuronal oscillations. Here, we examined whether tACS targeting M1 immediately following explicit motor sequence training is capable of modulating motor skill consolidation in older adults. In two sets of double-blind, sham-controlled experiments, tACS targeting left M1 was applied at either 10 Hz (alpha-tACS) or 20 Hz (beta-tACS) immediately after termination of a motor sequence training with the right (dominant) hand. Task performance was retested after an interval of 6 hours to assess consolidation of the training-acquired skill. EEG was recorded over left M1 to be able to detect local after-effects on oscillatory activity induced by tACS. Relative to the sham intervention, consolidation was selectively disrupted by posttraining alpha-tACS of M1, while posttraining beta-tACS of M1 had no effect on delayed retest performance compared to the sham intervention. No significant postinterventional changes of oscillatory activity in M1 were detected following alpha-tACS or beta-tACS. Our findings point to a frequency-specific interaction of tACS with posttraining motor memory processing and may suggest an inhibitory role of immediate posttraining alpha oscillations in M1 with respect to motor consolidation in healthy older adults.

Background: Motor learning is impaired in Parkinson's disease (PD), with patients demonstrating deficits in skill acquisition (online learning) and consolidation (offline learning) compared to healthy adults of similar age. Recent studies in young adults suggest that single bouts of aerobic exercise (AEX), performed in close temporal proximity to practicing a new motor task, may facilitate motor skill learning. Thus, we aimed at investigating the effects of a single bout of aerobic cycling on online and offline learning in PD patients. Methods: 17 PD patients (Hoehn and Yahr 1 - 2.5, age: 64.4 ± 6.2) participated in this crossover study. Immediately prior to practicing a novel balance task, patients either performed 30 min of (i) moderate intensity (60-70% VO2max) aerobic cycling, or (ii) seated rest (order counterbalanced). The task required patients to stabilize a balance platform (stabilometer) in a horizontal position for 30 s. For each experimental condition, patients performed 15 acquisition trials, followed by a retention test 24 h later. We calculated time in balance (platform within ± 5° from horizontal) for each trial, and analyzed within- and between-subjects differences in skill acquisition (online learning) and skill retention (offline learning) using mixed repeated-measures ANOVA. Results: We found that the exercise bout had no effect on performance level or online gains during acquisition, despite affecting the time course of skill improvements (larger initial and reduced late skill gains). Aerobic cycling significantly improved offline learning, as reflected by larger 24-h skill retention compared to the rest condition. Conclusion: Our results suggest that a single bout of moderate-intensity AEX is effective in improving motor skill consolidation in PD patients. Thus, acute exercise may represent an effective strategy to enhance motor memory formation in this population. More work is necessary to understand the underlying mechanisms, the optimal scheduling of exercise, and the applicability to other motor tasks. Further, the potential for patients in later disease stages need to be investigated. The study was a priori registered at ClinicalTrials.gov (NCT03245216).

THE BENEFICIAL ROLE OF SLEEP IN MOTOR MEMORY CONSOLIDATION IS NOW WELL DOCUMENTED. FOR EXAMPLE, AMPLE EVIDENCE INDICATES THAT consolidation of a newly learned sequence of movements, defined as a spontaneous (offline) improvement in performance that emerges in the absence of any further practice, is sleep dependent.1–3 Indeed, our group4 and others have demonstrated that delayed gains on a sequential finger-tapping task are triggered only after a night of sleep, whereas a comparable interval without sleep provides no additional benefit. Despite the accumulating evidence in support of sleep-dependent motor skill consolidation, there is still no consensus with respect to the sleep stages that are preferentially involved in this memory phase.5 Several studies support the hypothesis that post-training REM sleep is required for efficient motor skill consolidation.1,7–10 Yet other recent investigations have emphasized the importance of NREM sleep, including delta11 and stage 2 sleep.12 For example, Huber and colleagues11 have shown that slow wave activity (< 4 Hz) in the right parietal area during post-training sleep increases following practice on a visuomotor adaptation task, while others have reported that memory on a simple motor skill (i.e., the rotary pursuit task) is vulnerable to stage 2 sleep deprivation but not to REM sleep loss.13 Finally, Walker and colleagues3 demonstrated that improved overnight performance on a motor sequence learning task is associated with the amount of stage 2 sleep in the last quarter of the night. There has been increasing attention to the possible contribution of sleep spindles to memory processes.14 Spindles are an essential feature of stage 2 sleep that also appear throughout the depolarizing phase of slow wave sleep oscillation (