Intensity-dependent effects of tDCS on motor learning are related to dopamine.

Non-invasive brain stimulation techniques, such as transcranial direct current stimulation (tDCS), are popular methods for inducing neuroplastic changes to alter cognition and behaviour. One challenge for the field is to optimise stimulation protocols to maximise benefits. For this to happen, we need a better understanding of how stimulation modulates cortical functioning/behaviour. To date, there is increasing evidence for a dose-response relationship between tDCS and brain excitability, however how this relates to behaviour is not well understood. Even less is known about the neurochemical mechanisms which may drive the dose-response relationship between stimulation intensities and behaviour. Here, we examine the effect of three different tDCS stimulation intensities (1 mA, 2 mA, 4 mA anodal motor cortex tDCS) administered during the explicit learning of motor sequences. Further, to assess the role of dopamine in the dose-response relationship between tDCS intensities and behaviour, we examined how pharmacologically increasing dopamine availability, via 100 mg of levodopa, modulated the effect of stimulation on learning. In the absence of levodopa, we found that 4 mA tDCS improved and 1 mA tDCS impaired acquisition of motor sequences relative to sham stimulation. Conversely, levodopa reversed the beneficial effect of 4 mA tDCS. This effect of levodopa was no longer evident at the 48-h follow-up, consistent with previous work characterising the persistence of neuroplastic changes in the motor cortex resulting from combining levodopa with tDCS. These results provide the first direct evidence for a role of dopamine in the intensity-dependent effects of tDCS on behaviour.

Assessing the influence of dopamine and mindfulness on the formation of routines in visual search.

Given experience in cluttered but stable visual environments, our eye-movements form stereotyped routines that sample task-relevant locations, while not mixing-up routines between similar task-settings. Both dopamine signaling and mindfulness have been posited as factors that influence the formation of such routines, yet quantification of their impact remains to be tested in healthy humans. Over two sessions, participants searched through grids of doors to find hidden targets, using a gaze-contingent display. Within each session, door scenes appeared in either one of two colors, with each color signaling a differing set of likely target locations. We derived measures for how well target locations were learned (target-accuracy), how routine were sets of eye-movements (stereotypy), and the extent of interference between the two scenes (setting-accuracy). Participants completed two sessions, where they were administered either levodopa (dopamine precursor) or placebo (vitamin C), under double-blind counterbalanced conditions. Dopamine and trait mindfulness (assessed by questionnaire) interacted to influence both target-accuracy and stereotypy. Increasing dopamine improved accuracy and reduced stereotypy for high mindfulness scorers, but induced the opposite pattern for low mindfulness scorers. Dopamine also disrupted setting-accuracy invariant to mindfulness. Our findings show that mindfulness modulates the impact of dopamine on the target-accuracy and stereotypy of eye-movement routines, whereas increasing dopamine promotes interference between task-settings, regardless of mindfulness. These findings provide a link between non-human and human models regarding the influence of dopamine on the formation of task-relevant eye-movement routines and provide novel insights into behavior-trait factors that modulate the use of experience when building adaptive repertoires.

Exploring the intra-individual reliability of tDCS: A registered report.

Transcranial direct current stimulation (tDCS), a form of non-invasive brain stimulation, has become an important tool for the study of in-vivo brain function due to its modulatory effects. Over the past two decades, interest in the influence of tDCS on behaviour has increased markedly, resulting in a large body of literature spanning multiple domains. However, the effect of tDCS on human performance often varies, bringing into question the reliability of this approach. While reviews and meta-analyses highlight the contributions of methodological inconsistencies and individual differences, no published studies have directly tested the intra-individual reliability of tDCS effects on behaviour. Here, we conducted a large scale, double-blinded, sham-controlled registered report to assess the reliability of two single-session low-dose tDCS montages, previously found to impact response selection and motor learning operations, across two separate time periods. Our planned analysis found no evidence for either protocol being effective nor reliable. Post-hoc explorative analyses found evidence that tDCS influenced motor learning, but not response selection learning. In addition, the reliability of motor learning performance across trials was shown to be disrupted by tDCS. These findings are amongst the first to shed light specifically on the intra-individual reliability of tDCS effects on behaviour and provide valuable information to the field.

Dopamine Increases Accuracy and Lengthens Deliberation Time in Explicit Motor Skill Learning.

Although animal research implicates a central role for dopamine in motor skill learning, a direct causal link has yet to be established in neurotypical humans. Here, we tested if a pharmacological manipulation of dopamine alters motor learning, using a paradigm which engaged explicit, goal-directed strategies. Participants (27 females; 11 males; aged 18-29 years) first consumed either 100 mg of levodopa (n = 19), a dopamine precursor that increases dopamine availability, or placebo (n = 19). Then, during training, participants learnt the explicit strategy of aiming away from presented targets by instructed angles of varying sizes. Targets jumped mid-movement by the instructed aiming angle. Task success was thus contingent upon aiming accuracy and not speed. The effect of the dopamine manipulations on skill learning was assessed during training and after an overnight follow-up. Increasing dopamine availability at training improved aiming accuracy and lengthened reaction times, particularly for larger, more difficult aiming angles, both at training and, importantly, at follow-up, despite prominent session-by-session performance improvements in both accuracy and speed. Exogenous dopamine thus seems to result in a learnt, persistent propensity to better adhere to task goals. Results support the proposal that dopamine is important in engagement of instrumental motivation to optimize adherence to task goals, particularly when learning to execute goal-directed strategies in motor skill learning.

Individual Differences in Decision Strategy Relate to Neurochemical Excitability and Cortical Thickness.

The speed-accuracy trade-off (SAT), whereby faster decisions increase the likelihood of an error, reflects a cognitive strategy humans must engage in during the performance of almost all daily tasks. To date, computational modeling has implicated the latent decision variable of response caution (thresholds), the amount of evidence required for a decision to be made, in the SAT. Previous imaging has associated frontal regions, notably the left prefrontal cortex and the presupplementary motor area (pre-SMA), with the setting of such caution levels. In addition, causal brain stimulation studies, using transcranial direct current stimulation (tDCS), have indicated that while both of these regions are involved in the SAT, their role appears to be dissociable. tDCS efficacy to impact decision-making processes has previously been linked with neurochemical concentrations and cortical thickness of stimulated regions. However, to date, it is unknown whether these neurophysiological measures predict individual differences in the SAT, and brain stimulation effects on the SAT. Using ultra-high field (7T) imaging, here we report that instruction-based adjustments in caution are associated with both neurochemical excitability (the balance between GABA+ and glutamate) and cortical thickness across a range of frontal regions in both sexes. In addition, cortical thickness, but not neurochemical concentrations, was associated with the efficacy of left prefrontal and superior medial frontal cortex (SMFC) stimulation to modulate performance. Overall, our findings elucidate key neurophysiological predictors, frontal neural excitation, of individual differences in latent psychological processes and the efficacy of stimulation to modulate these.SIGNIFICANCE STATEMENT The speed-accuracy trade-off (SAT), faster decisions increase the likelihood of an error, reflects a cognitive strategy humans must engage in during most daily tasks. The SAT is often investigated by explicitly instructing participants to prioritize speed or accuracy when responding to stimuli. Using ultra-high field (7T) magnetic resonance imaging (MRI), we found that individual differences in the extent to which participants adjust their decision strategies with instruction related to neurochemical excitability (ratio of GABA+ to glutamate) and cortical thickness in the frontal cortex. Moreover, brain stimulation to the left prefrontal cortex and the superior medial frontal cortex (SMFC) modulated performance, with the efficacy specifically related to cortical thickness. This work sheds new light on the neurophysiological basis of decision strategies and brain stimulation.

Dopamine Alters the Effect of Brain Stimulation on Decision-Making.

Noninvasive brain stimulation techniques, such as transcranial direct current stimulation (tDCS), show promise in treating a range of psychiatric and neurologic conditions. However, optimization of such applications requires a better understanding of how tDCS alters cognition and behavior. Existing evidence implicates dopamine in tDCS alterations of brain activity and plasticity; however, there is as yet no causal evidence for a role of dopamine in tDCS effects on cognition and behavior. Here, in a preregistered, double-blinded study, we examined how pharmacologically manipulating dopamine altered the effect of tDCS on the speed-accuracy trade-off, which taps ubiquitous strategic operations. Cathodal tDCS was delivered over the left prefrontal cortex and the superior medial frontal cortex before participants (N = 62, 24 males, 38 females) completed a dot-motion task, making judgments on the direction of a field of moving dots under instructions to emphasize speed, accuracy, or both. We leveraged computational modeling to uncover how our interventions altered latent decisional processes driving the speed-accuracy trade-off. We show that dopamine in combination with tDCS (but not tDCS alone nor dopamine alone) not only impaired decision accuracy but also impaired discriminability, which suggests that these manipulations altered the encoding or representation of discriminative evidence. This is, to the best of our knowledge, the first direct evidence implicating dopamine in the way tDCS affects cognition and behavior.SIGNIFICANCE STATEMENT tDCS can improve cognitive and behavioral impairments in clinical conditions; however, a better understanding of its mechanisms is required to optimize future clinical applications. Here, using a pharmacological approach to manipulate brain dopamine levels in healthy adults, we demonstrate a role for dopamine in the effects of tDCS in the speed-accuracy trade-off, a strategic cognitive process ubiquitous in many contexts. In doing so, we provide direct evidence implicating dopamine in the way tDCS affects cognition and behavior.

Using EEG to study sensorimotor adaptation.

Sensorimotor adaptation, or the capacity to flexibly adapt movements to changes in the body or the environment, is crucial to our ability to move efficiently in a dynamic world. The field of sensorimotor adaptation is replete with rigorous behavioural and computational methods, which support strong conceptual frameworks. An increasing number of studies have combined these methods with electroencephalography (EEG) to unveil insights into the neural mechanisms of adaptation. We review these studies: discussing EEG markers of adaptation in the frequency and the temporal domain, EEG predictors for successful adaptation and how EEG can be used to unmask latent processes resulting from adaptation, such as the modulation of spatial attention. With its high temporal resolution, EEG can be further exploited to deepen our understanding of sensorimotor adaptation.

The acute effects of aerobic exercise on sensorimotor adaptation in chronic stroke.

BACKGROUND: Sensorimotor adaptation, or the capacity to adapt movement to changes in the moving body or environment, is a form of motor learning that is important for functional independence (e.g., regaining stability after slips or trips). Aerobic exercise can acutely improve many forms of motor learning in healthy adults. It is not known, however, whether acute aerobic exercise has similar positive effects on sensorimotor adaptation in stroke survivors as it does in healthy individuals. OBJECTIVE: The aim of this study was to determine whether acute aerobic exercise promotes sensorimotor adaptation in people post stroke. METHODS: A single-blinded crossover study. Participants attended two separate sessions, completing an aerobic exercise intervention in one session and a resting control condition in the other session. Sensorimotor adaptation was assessed before and after each session, as was brain derived neurotrophic factor. Twenty participants with chronic stroke completed treadmill exercise at moderate to high intensity for 30 minutes. RESULTS: Acute aerobic exercise in chronic stroke survivors significantly increased sensorimotor adaptation from pre to post treadmill intervention. The 30-minute treadmill intervention resulted in an averaged 2.99 ng/ml increase in BDNF levels (BDNF pre-treadmill = 22.31 + /-2.85 ng/ml, post-treadmill was = 25.31 + /-2.46 pg/ml; t(16) = 2.146, p = 0.048, cohen's d = 0.521, moderate effect size). CONCLUSIONS: These results indicate a potential role for aerobic exercise to promote the recovery of sensorimotor function in chronic stroke survivors.

How groove in music affects gait.

Rhythmic auditory stimulation (RAS) is a gait intervention in which gait-disordered patients synchronise footsteps to music or metronome cues. Musical 'groove', the tendency of music to induce movement, has previously been shown to be associated with faster gait, however, why groove affects gait remains unclear. One mechanism by which groove may affect gait is that of beat salience: music that is higher in groove has more salient musical beats, and higher beat salience might reduce the cognitive demands of perceiving the beat and synchronizing footsteps to it. If groove's effects on gait are driven primarily by the impact of beat salience on cognitive demands, then groove's effects might only be present in contexts in which it is relevant to reduce cognitive demands. Such contexts could include task parameters that increase cognitive demands (such as the requirement to synchronise to the beat), or individual differences that may make synchronisation more cognitively demanding. Here, we examined whether high beat salience can account for the effects of high-groove music on gait. First, we increased the beat salience of low-groove music to be similar to that of high-groove music by embedding metronome beats in low and high-groove music. We examined whether low-groove music with high beat salience elicited similar effects on gait as high-groove music. Second, we examined the effect of removing the requirement to synchronise footsteps to the beat (i.e., allowing participants to walk freely with the music), which is thought to remove the cognitive demand of synchronizing movements to the beat. We tested two populations thought to be sensitive to the cognitive demands of synchronisation, weak beat-perceivers and older adults. We found that increasing the beat salience of low-groove music increased stride velocity, but strides were still slower than with high-groove music. Similarly, removing the requirement to synchronise elicited faster, less variable gait, and reduced bias for stability, but high-groove music still elicited faster strides than low-groove music. These findings suggest that beat salience contributes to groove's effect on gait, but it does not fully account for it. Despite reducing task difficulty by equalizing beat salience and removing the requirement to synchronise, high-groove music still elicited faster, less variable gait. Therefore, other properties of groove also appear to play a role in groove's effect on gait.

Acoustic stimulation increases implicit adaptation in sensorimotor adaptation.

Sensorimotor adaptation is an important part of our ability to perform novel motor tasks (i.e., learning of motor skills). Efforts to improve adaptation in healthy and clinical patients using non-invasive brain stimulation methods have been hindered by inter-individual and intra-individual variability in brain susceptibility to stimulation. Here, we explore unpredictable loud acoustic stimulation as an alternative method of modulating brain excitability to improve sensorimotor adaptation. In two experiments, participants moved a cursor towards targets, and adapted to a 30º rotation of cursor feedback, either with or without unpredictable acoustic stimulation. Acoustic stimulation improved initial adaptation to sensory prediction errors in Study 1, and improved overnight retention of adaptation in Study 2. Unpredictable loud acoustic stimulation might thus be a potent method of modulating sensorimotor adaptation in healthy adults.

Task Feedback Processing Differs Between Young and Older Adults in Visuomotor Rotation Learning Despite Similar Initial Adaptation and Savings.

Ageing has been suggested to affect sensorimotor adaptation by impairing explicit strategy use. Here we recorded electrophysiological (EEG) responses during visuomotor rotation in both young (n = 24) and older adults (n = 25), to investigate the neural processes that underpin putative age-related effects on adaptation. We measured the feedback related negativity (FRN) and the P3 in response to task-feedback, as electrophysiological markers of task error processing and outcome evaluation. The two age groups adapted similarly and showed comparable after effects and savings when re-exposed to the same perturbation several days after the initial session. Older adults, however, had less distinct EEG responses (i.e., reduced FRN amplitudes) to negative and positive task feedback. The P3 did not differ between age groups. Both young and older adults also showed a sustained late positivity following task feedback. Measured at the frontal electrode Fz, this sustained activity was negatively associated with both the amount of voluntary disengagement of explicit strategy and savings. In conclusion, despite preserved task performance, we find clear differences in neural responses to errors in older people, which suggests that there is a fundamental decline in this aspect of sensorimotor brain function with age.

Task Errors Drive Memories That Improve Sensorimotor Adaptation.

Traditional views of sensorimotor adaptation (i.e., adaptation of movements to perturbed sensory feedback) emphasize the role of automatic, implicit correction of sensory prediction errors. However, latent memories formed during sensorimotor adaptation, manifest as improved relearning (e.g., savings), have recently been attributed to strategic corrections of task errors (failures to achieve task goals). To dissociate contributions of task errors and sensory prediction errors to latent sensorimotor memories, we perturbed target locations to remove or enforce task errors during learning and/or test, with male/female human participants. Adaptation improved after learning in all conditions where participants were permitted to correct task errors, and did not improve whenever we prevented correction of task errors. Thus, previous correction of task errors was both necessary and sufficient to improve adaptation. In contrast, a history of sensory prediction errors was neither sufficient nor obligatory for improved adaptation. Limiting movement preparation time showed that the latent memories driven by learning to correct task errors take at least two forms: a time-consuming but flexible component, and a rapidly expressible, inflexible component. The results provide strong support for the idea that movement corrections driven by a failure to successfully achieve movement goals underpin motor memories that manifest as savings. Such persistent memories are not exclusively mediated by time-consuming strategic processes but also comprise a rapidly expressible but inflexible component. The distinct characteristics of these putative processes suggest dissociable underlying mechanisms, and imply that identification of the neural basis for adaptation and savings will require methods that allow such dissociations.SIGNIFICANCE STATEMENT Latent motor memories formed during sensorimotor adaptation manifest as improved adaptation when sensorimotor perturbations are reencountered. Conflicting theories suggest that this "savings" is underpinned by different mechanisms, including a memory of successful actions, a memory of errors, or an aiming strategy to correct task errors. Here we show that learning to correct task errors is sufficient to show improved subsequent adaptation with respect to naive performance, even when tested in the absence of task errors. In contrast, a history of sensory prediction errors is neither sufficient nor obligatory for improved adaptation. Finally, we show that latent sensorimotor memories driven by task errors comprise at least two distinct components: a time-consuming, flexible component, and a rapidly expressible, inflexible component.

Task errors contribute to implicit aftereffects in sensorimotor adaptation.

Perturbations of sensory feedback evoke sensory prediction errors (discrepancies between predicted and actual sensory outcomes of movements), and reward prediction errors (discrepancies between predicted rewards and actual rewards). When our task is to hit a target, we expect to succeed in hitting the target, and so we experience a reward prediction error if the perturbation causes us to miss it. These discrepancies between intended task outcomes and actual task outcomes, termed "task errors," are thought to drive the use of strategic processes to restore success, although their role is incompletely understood. Here, as participants adapted to a 30° rotation of cursor feedback representing hand position, we investigated the role of task errors in sensorimotor adaptation: during target-reaching, we either removed task errors by moving the target mid-movement to align with cursor feedback of hand position, or enforced task error by moving the target away from the cursor feedback of hand position, by 20-30° randomly (clockwise in half the trials, counterclockwise in half the trials). Removing task errors not only reduced the extent of adaptation during exposure to the perturbation, but also reduced the amount of post-adaptation aftereffects that persisted despite explicit knowledge of the perturbation removal. Hence, task errors contribute to implicit adaptation resulting from sensory prediction errors. This suggests that the system which predicts the sensory consequences of actions via exposure to sensory prediction errors is also sensitive to reward prediction errors.

Triggering Mechanisms for Motor Actions: The Effects of Expectation on Reaction Times to Intense Acoustic Stimuli.

Motor actions can be released much sooner than normal when the go-signal is of very high intensity (>100 dBa). Although statistical evidence from individual studies has been mixed, it has been assumed that sternocleidomastoid (SCM) muscle activity could be used to distinguish between two neural circuits involved in movement triggering. We summarized meta-analytically the available evidence for this hypothesis, comparing the difference in premotor reaction time (RT) of actions where SCM activity was elicited (SCM+ trials) by loud acoustic stimuli against trials in which it was absent (SCM- trials). We found ten studies, all reporting comparisons between SCM+ and SCM- trials. Our mini meta-analysis showed that premotor RTs are faster in SCM+ than in SCM- trials, but the effect can be confounded by the variability of the foreperiods employed. We present experimental data showing that foreperiod predictability can induce differences in RT that would be of similar size to those attributed to the activation of different neurophysiological pathways to trigger prepared actions. We discuss plausible physiological mechanisms that would explain differences in premotor RTs between SCM+ and SCM- trials.

The role of attention and intention in synchronization to music: effects on gait.

Anecdotal accounts suggest that individuals spontaneously synchronize their movements to the 'beat' of background music, often without intending to, and perhaps even without attending to the music at all. However, the question of whether intention and attention are necessary to synchronize to the beat remains unclear. Here, we compared whether footsteps during overground walking were synchronized to the beat when young healthy adults were explicitly instructed to synchronize (intention to synchronize), and were not instructed to synchronize (no intention) (Experiment 1: intention). We also examined whether reducing participants' attention to the music affected synchronization, again when participants were explicitly instructed to synchronize, and when they were not (Experiment 2: attention/intention). Synchronization was much less frequent when no instructions to synchronize were given. Without explicit instructions to synchronize, there was no evidence of synchronization in 60% of the trials in Experiment 1, and 43% of the trials in Experiment 2. When instructed to synchronize, only 26% of trials in Experiment 1, and 14% of trials in Experiment 2 showed no evidence of synchronization. Because walking to music alters gait, we also examined how gait kinematics changed with or without instructions to synchronize, and attention to the music was required for synchronization to occur. Instructions to synchronize elicited slower, shorter, and more variable strides than walking in silence. Reducing attention to the music did not significantly affect synchronization of footsteps to the beat, but did elicit slower gait. Thus, during walking, intention, but not attention, appears to be necessary to synchronize footsteps to the beat, and synchronization elicits slower, shorter, and more variable strides, at least in young healthy adults.

Cerebellar anodal tDCS increases implicit learning when strategic re-aiming is suppressed in sensorimotor adaptation.

Neurophysiological and neuroimaging work suggests that the cerebellum is critically involved in sensorimotor adaptation. Changes in cerebellar function alter behaviour when compensating for sensorimotor perturbations, as shown by non-invasive stimulation of the cerebellum and studies involving patients with cerebellar degeneration. It is known, however, that behavioural responses to sensorimotor perturbations reflect both explicit processes (such as volitional aiming to one side of a target to counteract a rotation of visual feedback) and implicit, error-driven updating of sensorimotor maps. The contribution of the cerebellum to these explicit and implicit processes remains unclear. Here, we examined the role of the cerebellum in sensorimotor adaptation to a 30° rotation of visual feedback of hand position during target-reaching, when the capacity to use explicit processes was manipulated by controlling movement preparation times. Explicit re-aiming was suppressed in one condition by requiring subjects to initiate their movements within 300ms of target presentation, and permitted in another condition by requiring subjects to wait approximately 1050ms after target presentation before movement initiation. Similar to previous work, applying anodal transcranial direct current stimulation (tDCS; 1.5mA) to the right cerebellum during adaptation resulted in faster compensation for errors imposed by the rotation. After exposure to the rotation, we evaluated implicit remapping in no-feedback trials after providing participants with explicit knowledge that the rotation had been removed. Crucially, movements were more adapted in these no-feedback trials following cerebellar anodal tDCS than after sham stimulation in both long and short preparation groups. Thus, cerebellar anodal tDCS increased implicit remapping during sensorimotor adaptation, irrespective of preparation time constraints. The results are consistent with the possibility that the cerebellum contributes to the formation of new visuomotor maps that correct perturbations in sensory feedback, even when explicit processes are suppressed during sensorimotor adaptation.

Estimating the implicit component of visuomotor rotation learning by constraining movement preparation time.

When sensory feedback is perturbed, accurate movement is restored by a combination of implicit processes and deliberate reaiming to strategically compensate for errors. Here, we directly compare two methods used previously to dissociate implicit from explicit learning on a trial-by-trial basis: 1) asking participants to report the direction that they aim their movements, and contrasting this with the directions of the target and the movement that they actually produce, and 2) manipulating movement preparation time. By instructing participants to reaim without a sensory perturbation, we show that reaiming is possible even with the shortest possible preparation times, particularly when targets are narrowly distributed. Nonetheless, reaiming is effortful and comes at the cost of increased variability, so we tested whether constraining preparation time is sufficient to suppress strategic reaiming during adaptation to visuomotor rotation with a broad target distribution. The rate and extent of error reduction under preparation time constraints were similar to estimates of implicit learning obtained from self-report without time pressure, suggesting that participants chose not to apply a reaiming strategy to correct visual errors under time pressure. Surprisingly, participants who reported aiming directions showed less implicit learning according to an alternative measure, obtained during trials performed without visual feedback. This suggests that the process of reporting can affect the extent or persistence of implicit learning. The data extend existing evidence that restricting preparation time can suppress explicit reaiming and provide an estimate of implicit visuomotor rotation learning that does not require participants to report their aiming directions.NEW & NOTEWORTHY During sensorimotor adaptation, implicit error-driven learning can be isolated from explicit strategy-driven reaiming by subtracting self-reported aiming directions from movement directions, or by restricting movement preparation time. Here, we compared the two methods. Restricting preparation times did not eliminate reaiming but was sufficient to suppress reaiming during adaptation with widely distributed targets. The self-report method produced a discrepancy in implicit learning estimated by subtracting aiming directions and implicit learning measured in no-feedback trials.

Savings for visuomotor adaptation require prior history of error, not prior repetition of successful actions.

When we move, perturbations to our body or the environment can elicit discrepancies between predicted and actual outcomes. We readily adapt movements to compensate for such discrepancies, and the retention of this learning is evident as savings, or faster readaptation to a previously encountered perturbation. The mechanistic processes contributing to savings, or even the necessary conditions for savings, are not fully understood. One theory suggests that savings requires increased sensitivity to previously experienced errors: when perturbations evoke a sequence of correlated errors, we increase our sensitivity to the errors experienced, which subsequently improves error correction (Herzfeld et al. 2014). An alternative theory suggests that a memory of actions is necessary for savings: when an action becomes associated with successful target acquisition through repetition, that action is more rapidly retrieved at subsequent learning (Huang et al. 2011). In the present study, to better understand the necessary conditions for savings, we tested how savings is affected by prior experience of similar errors and prior repetition of the action required to eliminate errors using a factorial design. Prior experience of errors induced by a visuomotor rotation in the savings block was either prevented at initial learning by gradually removing an oppositely signed perturbation or enforced by abruptly removing the perturbation. Prior repetition of the action required to eliminate errors in the savings block was either deprived or enforced by manipulating target location in preceding trials. The data suggest that prior experience of errors is both necessary and sufficient for savings, whereas prior repetition of a successful action is neither necessary nor sufficient for savings.

Familiarity with music increases walking speed in rhythmic auditory cuing.

Rhythmic auditory stimulation (RAS) is a gait rehabilitation method in which patients synchronize footsteps to a metronome or musical beats. Although RAS with music can ameliorate gait abnormalities, outcomes vary, possibly because music properties, such as groove or familiarity, differ across interventions. To optimize future interventions, we assessed how initially familiar and unfamiliar low-groove and high-groove music affected synchronization accuracy and gait in healthy individuals. We also experimentally increased music familiarity using repeated exposure to initially unfamiliar songs. Overall, familiar music elicited faster stride velocity and less variable strides, as well as better synchronization performance (matching of step tempo to beat tempo). High-groove music, as reported previously, led to faster stride velocity than low-groove music. We propose two mechanisms for familiarity's effects. First, familiarity with the beat structure reduces cognitive demands of synchronizing, leading to better synchronization performance and faster, less variable gait. Second, familiarity might have elicited faster gait by increasing enjoyment of the music, as enjoyment was higher after repeated exposure to initially low-enjoyment songs. Future studies are necessary to dissociate the contribution of these mechanisms to the observed RAS effects of familiar music on gait.

Neural mechanisms of rhythm perception: present findings and future directions.

The capacity to synchronize movements to the beat in music is a complex, and apparently uniquely human characteristic. Synchronizing movements to the beat requires beat perception, which entails prediction of future beats in rhythmic sequences of temporal intervals. Absolute timing mechanisms, where patterns of temporal intervals are encoded as a series of absolute durations, cannot fully explain beat perception. Beat perception seems better accounted for by relative timing mechanisms, where temporal intervals of a pattern are coded relative to a periodic beat interval. Evidence from behavioral, neuroimaging, brain stimulation and neuronal cell recording studies suggests a functional dissociation between the neural substrates of absolute and relative timing. This chapter reviews current findings on relative timing in the context of rhythm and beat perception.