Decoding of spatial proportions using somatosensory feedback in sighted and visually impaired children.

BACKGROUND AND PURPOSE: Humans can naturally operate with ratios of continuous magnitudes (proportions). We asked if sighted children (S) and visually impaired children (VI) can discriminate proportions via somatosensory feedback. PROCEDURES: Children formed a proportion by tracing a pair of straight lines with their finger, and compared this proportion with a second proportion resulting from the tracing of another pair of lines. MAIN FINDINGS: Performance was 68% in S, thus significantly lower (p < 0.001) compared to VI (75%). Tracing velocity (p < 0.01) and trial-to-trial variability of tracing velocity (p < 0.05) was higher in S compared to VI. CONCLUSIONS: Operating with proportions solely from somatosensory feedback is possible, thus tracing lines might support learning in mathematics education. Kinematic variables point to the reason for the difference between S and VI, in that higher trial-to-trial variability in velocity in S leads to biased estimation of absolute line lengths.

Applying augmented feedback in basketball training facilitates improvements in jumping performance.

Augmented feedback, which is feedback about movement characteristics provided by an external source, can facilitate performance improvements. Results from recent studies, in which information about the jumping height were presented, indicated increased jumping performance due to augmented feedback. In the present study we aimed to utilize augmented feedback about the jumping height in regular basketball training. Therefore, augmented feedback was implemented and part of the training regime, and information about the jumping height was displayed while subjects performed catch and shoot exercises with the basketball. 18 subjects (9 with augmented feedback, 9 without augmented feedback) practiced for 3 weeks (9 training sessions). The subjects receiving augmented feedback, but not subjects in the control group, displayed increased jumping heights of countermovement-jumps and drop-jumps after the training. The ground contact times of countermovement-jumps and drop-jumps were not significantly changed after training. The number of successful hits, indicating ball performance, did also not change after training. Thus augmented feedback had no detrimental effect on sport-specific performance. The unchanged ground contact times of drop-jumps in combination with increased jumping heights indicate increased efficiency of stretch shortening cycle contractions. According to the positive effect of augmented feedback on jumping performance and the simplicity with which it was integrated into the training regime we recommend this method for regular basketball training.HighlightsAugmented feedback about the jumping height in basketball training facilitated jumping height of countermovement-jumps and drop-jumps.The ground contact times were not altered, suggesting increased efficiency of the stretch-shortening cycle during jumping.According to the positive effect and its simple implementation augmented feedback is recommended for regular basketball training.

Do Primary School Children Benefit from Drop-Jump Training with Different Schedules of Augmented Feedback about the Jump Height?

In children, the training of jumps leads to improved jumping and running performance. Augmented feedback about the jump height is known to facilitate performance improvements in adults. In the present study, the impact of augmented feedback on jumping performance was investigated in 4th grade primary school children executing drop-jump training for 8 weeks (24 sessions, 3 times/week). Ten children (eight males, two females, aged 9.6 ± 0.3 years), received feedback for 8 weeks, and 11 children (nine males, two females, aged 9.5 ± 0.2 years) received feedback only during the last 4 weeks. Drop-jumps training was integrated in physical education classes. Drop-jump and countermovement-jump heights were improved after 24 training sessions (p < 0.01 for both types of jumps in both groups). Ground contact times of drop-jumps were quite long (>200 ms) and not altered by training, and the reactive strength index of drop-jumps was between 0.75 and 1.5 in most children. Augmented feedback did not facilitate jumping performance like in previous studies with adult participants. In contrast, withholding augmented feedback during the first 4 weeks of training was associated with a reduction in jumping performance (p < 0.01 for drop-jumps, p < 0.05 for countermovement-jumps). Finally, improvements did not transfer to functional motor tasks containing jumps. According to the costs and outcomes we do not recommend drop-jump training with augmented feedback about the jump height for 4th grade physical education classes.

Determining the types of descending waves from transcranial magnetic stimulation measured with conditioned H-reflexes in humans.

Non-invasive techniques are scarce with which human (motor) cortical mechanisms can be investigated. In a series of previous experiments, we have applied an advanced form of conditioning technique with transcranial magnetic stimulation (TMS) and peripheral nerve stimulation by which excitability changes at the laminar level in the primary motor cortex can be estimated. This method builds on the assumption that the first of subsequent corticospinal waves from TMS which is assessed with H-reflexes (called early facilitation) results from indirect excitation of corticospinal neurons in motor cortex (I-wave) and not direct excitation of corticospinal axons (D-wave). So far, we have not provided strong experimental evidence that this is actually the case. In the present study, we therefore compared temporal differences of the early facilitation between transcranial magnetic and electrical stimulation (TES). TES is known to excite the axons of corticospinal neurons. TES in our study caused a temporal shift of the early facilitation of H-reflexes in all subjects compared to TMS, which indicates that the early facilitation with TMS is indeed produced by an I-wave. Additionally, we investigated temporal shifts of the early facilitation with different TMS intensities and two TMS coils. It has long been known that TMS with higher intensities can induce a D-wave. Accordingly, we found that TMS with an intensity of 150% of resting motor threshold compared to 130%/110% results in a temporal shift of the early facilitation, indicating the presence of a D-wave. This effect was dependent on the coil type.

Brain activation patterns during visuomotor adaptation in motor experts and novices: An FDG PET study with unrestricted movements.

BACKGROUND: Speed of performance improvements and the strength of memory consolidation in humans vary with movement expertise. Underlying neural mechanisms of behavioural differences between levels of movement expertise are so far unknown. NEW METHOD: In this study, PET with [18F]fluorodeoxyglucose (FDG) was proposed as a powerful novel methodology to assess learning-related brain activity patterns during large non-restricted movements (ball throwing with a right hand). 24 male handball players ('Experts') and 24 male participants without handball experience ('Novices') performed visuomotor adaptations to prismatic glasses with or without strategic manoeuvres (i.e., explicit or implicit adaptation). RESULTS: Regional changes in FDG uptake as a marker of neuronal activity, relative to a control condition, were assessed. Prismatic adaptation, in general, was associated with decreased occipital neuronal activity as a possible response to misleading visual information. In 'Experts', the adaptation was associated with altered neuronal activity in a network comprising the right parietal cortex and the left cerebellum. In 'Novices', implicit adaptation resulted in an activation of the middle frontal and inferior temporal gyrus. COMPARISON WITH EXISTING METHODS: This study demonstrates the versatility of FDG PET for studying brain activations patterns in experimental settings with unrestricted movements that are not accessible by other techniques (e.g., fMRI or EEG). CONCLUSIONS: Observed results are consistent with the involvement of different functional networks related to strategic manoeuvres and expertise levels. This strengthens the assumption of different mechanisms underlying behavioural changes associated with movement expertise. Furthermore, the present study underscores the value of FDG PET for studying brain activation patterns during unrestricted movements.

Balance Training Reduces Postural Sway and Improves Sport-specific Performance in Visually Impaired Cross-Country Skiers.

ABSTRACT: Kurz, A, Lauber, B, Franke, S, and Leukel, C. Balance training reduces postural sway and improves sport-specific performance in visually impaired cross-country skiers. J Strength Cond Res 35(1): 247-252, 2021-Balance training is highly effective in reducing sport injuries and causes improvements in postural stability and rapid force production. So far, the positive effects of balance training have been described for healthy athletes. In the present experiments, we questioned whether athletes with disabilities of the visual system can also benefit from balance training. Fourteen visually impaired cross-country skiers participated in this randomized controlled study. The intervention group (N = 7) completed 8 sessions of balance training over a period of 4 weeks (2 times per week), whereas a waiting control group (N = 7) received no training during that time. After training, postural sway was significantly reduced in the intervention group but not in the waiting control group. In addition, sport-specific performance, which was assessed by a standardized Cooper's 12-minute test on roller skis or rollerblades, increased in the intervention group. The change in postural sway from the premeasurement to the postmeasurement correlated with the change in sport-specific performance in all subjects. Our results indicate that balance training is useful for improving postural stability and sport-specific performance in visually impaired cross-country skiers. We propose that balance training should therefore be implemented as part of the training routine in athletes with disabilities of the visual system.

Training of a discrete motor skill in humans is accompanied by increased excitability of the fastest corticospinal connections at movement onset.

KEY POINTS: The primary motor cortex (M1) is fundamentally important for the acquisition of skilled motor behaviours. We tested the excitability changes of distinct M1 circuits at movement onset with TMS H-reflex conditioning. Human subjects trained a discrete spatiotemporal motor skill. Practice was associated with reduced kinematic variability and improved motor performance. Performance improvements were paralleled by task-specific excitability increases of the fastest corticospinal connections at infragranular layer 5b of M1. No task-related changes in excitability were observed at supragranular layers. Excitability changes in the fastest corticospinal connections were not directly related to changes in motor performance. ABSTRACT: The primary motor cortex (M1) is fundamentally important for the acquisition of skilled motor behaviours. Recent advances in imaging and electrophysiological techniques have improved our understanding of M1 neural circuit modulation in rodents and non-human primates during motor learning. However, little remains known about the learning-related changes of distinct elements in the human brain. In this study, we tested excitability changes of different neural circuits (infragranular and supragranular layers) in the M1 of human subjects who underwent training in a discrete spatiotemporal motor skill. Excitability modulations were assessed by recording H-reflex facilitation from transcranial magnetic stimulation at movement onset. Motor practice improved the consistency of movements and was accompanied by an excitability increase of the fastest corticospinal connections during the initial stages of motor practice. No such excitability changes were observed for training in a simple motor skill and circuits at supragranular layers of M1. Notably, changes in excitability were not associated with changes in motor performance. Our findings could reflect learning-related increases in the recruitment and/or reorganisation of the fastest corticospinal connections.

Evidence that distinct human primary motor cortex circuits control discrete and rhythmic movements.

KEY POINTS: Discrete and rhythmic dynamics are inherent components of (human) movements. We provide evidence that distinct human motor cortex circuits contribute to discrete and rhythmic movements. Excitability of supragranular layer circuits of the human motor cortex was higher during discrete movements than during rhythmic movements. Conversely, more complex corticospinal circuits showed higher excitability during rhythmic movements than during discrete movements. No task-specific differences existed for corticospinal output neurons at infragranular layers. The excitability differences were found to be time(phase)-specific and could not be explained by the kinematic properties of the movements. The same task-specific differences were found between the last cycle of a rhythmic movement period and ongoing rhythmic movements. ABSTRACT: Human actions entail discrete and rhythmic movements (DM and RM, respectively). Recent insights from human and animal studies indicate different neural control mechanisms for DM and RM, emphasizing the intrinsic nature of the task. However, how distinct human motor cortex circuits contribute to these movements remains largely unknown. In the present study, we tested distinct primary motor cortex and corticospinal circuits and proposed that they show differential excitability between DM and RM. Human subjects performed either 1) DM or 2) RM using their right wrist. We applied an advanced electrophysiological approach involving transcranial magnetic stimulation and peripheral nerve stimulation to test the excitability of the neural circuits. Probing was performed at different movement phases: movement initiation (MI, 20 ms after EMG onset) and movement execution (ME, 200 ms after EMG onset) of the wrist flexion. At MI, excitability at supragranular layers was significantly higher in DM than in RM. Conversely, excitability of more complex corticospinal circuits was significantly lower in DM than RM at ME. No task-specific differences were found for direct corticospinal output neurons at infragranular layers. The neural differences could not be explained by the kinematic properties of the movements and also existed between ongoing RM and the last cycle of RM. Our results therefore strengthen the hypothesis that different neural control mechanisms engage in DM and RM.

Excitability of Upper Layer Circuits Relates to Torque Output in Humans.

The relation between primary motor cortex (M1) activity and (muscular) force output has been studied extensively. Results from previous studies indicate that activity of a part of yet unidentified neurons in M1 are positively correlated with increased force levels. One considerable candidate causing this positive correlation could be circuits at supragranular layers. Here we tested this hypothesis and used the combination of H-reflexes with transcranial magnetic stimulation (TMS) to investigate laminar associations with force output in human subjects. Excitability of different M1 circuits were probed at movement onset and at peak torque while participants performed auxotonic contractions of the wrist with different torque levels. Only at peak torque we found a significant positive correlation between excitability of M1 circuits most likely involving neurons at supragranular layers and joint torque level. We argue that this finding may relate to the special role of upper layer circuits in integrating (force-related) afferent feedback and their connectivity with task-relevant pyramidal and also extrapyramidal pathways projecting to motoneurones in the spinal cord.

Non-invasive assessment of superficial and deep layer circuits in human motor cortex.

KEY POINTS: The first indirect (I) corticospinal volley from stimulation of the motor cortex consists of two parts: one that originates from infragranular layer 5 and a subsequent part with a delay of 0.6 ms to which supragranular layers contribute. Non-invasive probing of these two parts was performed in humans using a refined electrophysiological method involving transcranial magnetic stimulation and peripheral nerve stimulation. Activity modulation of these two parts during a sensorimotor discrimination task was consistent with previous results in monkeys obtained with laminar recordings. ABSTRACT: Circuits in superficial and deep layers play distinct roles in cortical computation, but current methods to study them in humans are limited. Here, we developed a novel approach for non-invasive assessment of layer-specific activity in the human motor cortex. We first conducted brain slice and in vivo experiments on monkey motor cortex to investigate the output timing from layer 5 (including corticospinal neurons) following extracellular stimulation. Neuron responses contained cyclical waves. The first wave was composed of two parts: the earliest part originated only from stimulation of layer 5; after 0.6 ms, stimuli to superficial layers 2/3 could also contribute. In healthy humans we then assessed different parts of the first corticospinal volley elicited by transcranial magnetic stimulation (TMS), by interacting TMS with stimulation of the median nerve generating an H-reflex. By adjusting the delay between stimuli, we could assess the earliest volley evoked by TMS, and the part 0.6 ms later. Measurements were made while subjects performed a visuo-motor discrimination task, which has been previously shown in monkey to modulate superficial motor cortical cells selectively depending on task difficulty. We showed a similar selective modulation of the later part of the TMS volley, as expected if this part of the volley is sensitive to superficial cortical excitability. We conclude that it is possible to segregate different cortical circuits which may refer to different motor cortex layers in humans, by exploiting small time differences in the corticospinal volleys evoked by non-invasive stimulation.

Evidence for a subcortical contribution to intracortical facilitation.

Intracortical facilitation (ICF) describes the facilitation of an EMG response (motor evoked potential) to a suprathreshold pulse (S2) of transcranial magnetic stimulation (TMS) by a preceding subthreshold pulse (S1) given 10-15 ms earlier. ICF is widely assumed to originate from intracortical mechanisms. In this study, we used spinal H-reflexes to test whether subcortical mechanisms can also contribute to the facilitation. Measurements were performed in the upper limb muscle flexor carpi radialis in 17 healthy volunteers, and in the lower limb muscle soleus in 16 healthy volunteers. S2 given alone facilitated the H-reflex. When S1 preceded S2 by 10 ms, the amount of facilitation increased, compatible with ICF. However, S1 given alone also facilitated the H-reflex, suggesting that it had evoked descending activity even though its intensity was well below resting motor threshold. Across participants, the amount of H-reflex facilitation from S1 alone was proportional to the degree of H-reflex facilitation with combined S1-S2. These results indicate that subcortical mechanisms can contribute to ICF and potentially add to the variability of the ICF measure reported in previous studies.

Assessing TMS-induced D and I waves with spinal H-reflexes.

Transcranial magnetic stimulation (TMS) of motor cortex produces a series of descending volleys known as D (direct) and I (indirect) waves. In the present study, we questioned whether spinal H-reflexes can be used to dissect D waves and early and late I waves from TMS. We therefore probed H-reflex facilitation at arrival times of D and I waves at the spinal level and thereby changed TMS parameters that have previously been shown to have selective effects on evoked D and different I waves. We changed TMS intensity and current direction and applied a double-pulse paradigm known as short-interval intracortical inhibition (SICI). Experiments were conducted in flexor carpi radialis (FCR) in the arm and soleus (SOL) in the leg. There were two major findings: 1) in FCR, H-reflex facilitation showed characteristic modulations with altered TMS parameters that correspond to the changes of evoked D and I waves; and 2) H-reflexes in SOL did not, possibly because of increased interference from other spinal circuits. Therefore, the most significant outcome of this study is that in FCR, H-reflexes combined with TMS seem to be a useful technique to dissect TMS-induced D and I waves. NEW & NOTEWORTHY Questions that relate to corticospinal function in pathophysiology and movement control demand sophisticated techniques to provide information about corticospinal mechanisms. We introduce a noninvasive electrophysiological technique that may be useful in describing such mechanisms in more detail by dissecting D and I waves from transcranial magnetic stimulation (TMS). Based on the combination of spinal H-reflexes and TMS in the flexor carpi radialis muscle, the technique was shown to measure selective effects on D and I waves from changing TMS parameters.

Non-invasive Assessment of Changes in Corticomotoneuronal Transmission in Humans.

The corticospinal pathway is the major pathway connecting the brain with the muscles and is therefore highly relevant for movement control and motor learning. There exists a number of noninvasive electrophysiological methods investigating the excitability and plasticity of this pathway. However, most methods are based on quantification of compound potentials and neglect that the corticospinal pathway consists of many different connections that are more or less direct. Here, we present a method that allows testing excitability of different fractions of the corticospinal transmission. This so called H-reflex conditioning technique allows one to assess excitability of the fastest (monosynaptic) and also polysynaptic corticospinal pathways. Furthermore, by using two different stimulation sites, the motor cortex and the cervicomedullary junction, it allows not only differentiation between cortical and spinal effects but also assessment of transmission at the corticomotoneural synapse. In this manuscript, we describe how this method can be used to assess corticomotoneural transmission after low-frequency repetitive transcranial magnetic stimulation, a method that was previously shown to reduce excitability of cortical cells. Here we demonstrate that not only cortical cells are affected by this repetitive stimulation but also transmission at the corticomotoneuronal synapse at the spinal level. This finding is important for the understanding of basic mechanisms and sites of neuroplasticity. Besides investigation of basic mechanisms, the H-reflex conditioning technique may be applied to test changes in corticospinal transmission following behavioral (e.g., training) or therapeutic interventions, pathology or aging and therefore allows a better understanding of neural processes that underlie movement control and motor learning.

Do placebo expectations influence perceived exertion during physical exercise?

This study investigates the role of placebo expectations in individuals' perception of exertion during acute physical exercise. Building upon findings from placebo and marketing research, we examined how perceived exertion is affected by expectations regarding a) the effects of exercise and b) the effects of the exercise product worn during the exercise. We also investigated whether these effects are moderated by physical self-concept. Seventy-eight participants conducted a moderate 30 min cycling exercise on an ergometer, with perceived exertion (RPE) measured every 5 minutes. Beforehand, each participant was randomly assigned to 1 of 4 conditions and watched a corresponding film clip presenting "scientific evidence" that the exercise would or would not result in health benefits and that the exercise product they were wearing (compression garment) would additionally enhance exercise benefits or would only be worn for control purposes. Participants' physical self-concept was assessed via questionnaire. Results partially demonstrated that participants with more positive expectations experienced reduced perceived exertion during the exercise. Furthermore, our results indicate a moderator effect of physical self-concept: Individuals with a high physical self-concept benefited (in terms of reduced perceived exertion levels) in particular from an induction of generally positive expectations. In contrast, individuals with a low physical self-concept benefited when positive expectations were related to the exercise product they were wearing. In sum, these results suggest that placebo expectations may be a further, previously neglected class of psychological factors that influence the perception of exertion.

Expectations affect psychological and neurophysiological benefits even after a single bout of exercise.

The study investigated whether typical psychological, physiological, and neurophysiological changes from a single exercise are affected by one's beliefs and expectations. Seventy-six participants were randomly assigned to four groups and saw different multimedia presentations suggesting that the subsequent exercise (moderate 30 min cycling) would result in more or less health benefits (induced expectations). Additionally, we assessed habitual expectations reflecting previous experience and beliefs regarding exercise benefits. Participants with more positive habitual expectations consistently demonstrated both greater psychological benefits (more enjoyment, mood increase, and anxiety reduction) and greater increase of alpha-2 power, assessed with electroencephalography. Manipulating participants' expectations also resulted in largely greater increases of alpha-2 power, but not in more psychological exercise benefits. On the physiological level, participants decreased their blood pressure after exercising, but this was independent of their expectations. These results indicate that habitual expectations in particular affect exercise-induced psychological and neurophysiological changes in a self-fulfilling manner.

Motor Experts Care about Consistency and Are Reluctant to Change Motor Outcome.

Thousands of hours of physical practice substantially change the way movements are performed. The mechanisms underlying altered behavior in highly-trained individuals are so far little understood. We studied experts (handballers) and untrained individuals (novices) in visuomotor adaptation of free throws, where subjects had to adapt their throwing direction to a visual displacement induced by prismatic glasses. Before visual displacement, experts expressed lower variability of motor errors than novices. Experts adapted and de-adapted slower, and also forgot the adaptation slower than novices. The variability during baseline was correlated with the learning rate during adaptation. Subjects adapted faster when variability was higher. Our results indicate that experts produced higher consistency of motor outcome. They were still susceptible to the sensory feedback informing about motor error, but made smaller adjustments than novices. The findings of our study relate to previous investigations emphasizing the importance of action exploration, expressed in terms of outcome variability, to facilitate learning.

Force and Position Control in Humans - The Role of Augmented Feedback.

During motor behaviour, humans interact with the environment by for example manipulating objects and this is only possible because sensory feedback is constantly integrated into the central nervous system and these sensory inputs need to be weighted in order meet the task specific goals. Additional feedback presented as augmented feedback was shown to have an impact on motor control and motor learning. A number of studies investigated whether force or position feedback has an influence on motor control and neural activation. However, as in the previous studies the presentation of the force and position feedback was always identical, a recent study assessed whether not only the content but also the interpretation of the feedback has an influence on the time to fatigue of a sustained submaximal contraction and the (inhibitory) activity of the primary motor cortex using subthreshold transcranial magnetic stimulation. This paper describes one possible way to investigate the influence of the interpretation of feedback on motor behaviour by investigating the time to fatigue of submaximal sustained contractions together with the neuromuscular adaptations that can be investigated using surface EMG. Furthermore, the current protocol also describes how motor cortical (inhibitory) activity can be investigated using subthreshold TMS, a method known to act solely on the cortical level. The results show that when participants interpret the feedback as position feedback, they display a significantly shorter time to fatigue of a submaximal sustained contraction. Furthermore, subjects also displayed an increased inhibitory activity of the primary cortex when they believed to receive position feedback compared when they believed to receive force feedback. Accordingly, the results show that interpretation of feedback results in differences on a behavioural level (time to fatigue) that is also reflected in interpretation-specific differences in the amount of inhibitory M1 activity.

In Experts, underlying processes that drive visuomotor adaptation are different than in Novices.

Processes responsible for improvements in motor performance are often contrasted in an explicit and an implicit part. Explicit learning enables task success by using strategic (declarative) knowledge. Implicit learning refers to a change in motor performance without conscious effort. In this study, we tested the contribution of explicit and implicit processes in a visuomotor adaptation task in subjects with different expertise in the task they were asked to adapt. Thirty handball players (Experts) and 30 subjects without handball experience (Novices) participated. Three experiments tested visuomotor adaptation of a free throw in team handball using prismatic glasses. The difference between experiments was that in Experiment 2 and 3, contribution of explicit processes was prevented, whereas Experiment 1 allowed contribution of explicit and implicit processes. Retention was assessed in Experiment 3. There were three main findings: (i) contribution of explicit processes to adaptation was stronger in Experts than Novices (Experiment 1); (ii) adaptation took longer in Experts when preventing contribution of explicit processes (Experiment 2); and (iii) retention was stronger in Experts (Experiment 3). This study shows that learning processes involved in visuomotor adaptation change by expertise, with more involvement of explicit processes and most likely other implicit processes to adaptation in Experts.

Brain activity during observation and motor imagery of different balance tasks: an fMRI study.

After immobilization, patients show impaired postural control and increased risk of falling. Therefore, loss of balance control should already be counteracted during immobilization. Previously, studies have demonstrated that both motor imagery (MI) and action observation (AO) can improve motor performance. The current study elaborated how the brain is activated during imagination and observation of different postural tasks to provide recommendations about the conception of non-physical balance training. For this purpose, participants were tested in a within-subject design in an fMRI-scanner in three different conditions: (a) AO + MI, (b) AO, and (c) MI. In (a) participants were instructed to imagine themselves as the person pictured in the video whereas in (b) they were instructed simply to watch the video. In (c) subjects closed their eyes and kinesthetically imagined the task displayed in the video. Two tasks were evaluated in each condition: (i) static standing balance and (ii) dynamic standing balance (medio-lateral perturbation). In all conditions the start of a new trial was indicated every 2 sec by a sound. During AO + MI of the dynamic task, participants activated motor centers including the putamen, cerebellum, supplementary motor area, premotor cortices (PMv/d) and primary motor cortex (M1). MI showed a similar pattern but no activity in M1 and PMv/d. In the SMA and cerebellum, activity was generally higher in the dynamic than in the static condition. AO did not significantly activate any of these brain areas. Our results showed that (I) mainly AO + MI, but also MI, activate brain regions important for balance control; (II) participants display higher levels of brain activation in the more demanding balance task; (III) there is a significant difference between AO + MI and AO. Consequently, best training effects should be expected when participants apply MI during AO (AO + MI) of challenging postural tasks.

Repetitive activation of the corticospinal pathway by means of rTMS may reduce the efficiency of corticomotoneuronal synapses.

Low-frequency rTMS applied to the primary motor cortex (M1) may produce depression of motor-evoked potentials (MEPs). This depression is commonly assumed to reflect changes in cortical circuits. However, little is known about rTMS-induced effects on subcortical circuits. Therefore, the present study aimed to clarify whether rTMS influences corticospinal transmission by altering the efficiency of corticomotoneuronal (CM) synapses. The corticospinal transmission to soleus α-motoneurons was evaluated through conditioning of the soleus H-reflex by magnetic stimulation of either M1 (M1-conditioning) or the cervicomedullary junction (CMS-conditioning). The first facilitation of the H-reflex (early facilitation) was determined after M1- and CMS-conditioning. Comparison of the early facilitation before and after 20-min low-frequency (1 Hz) rTMS revealed suppression with M1- (-17 ± 4%; P = 0.001) and CMS-conditioning (-6 ± 2%; P = 0.04). The same rTMS protocol caused a significant depression of compound MEPs, whereas amplitudes of H-reflex and M-wave remained unaffected, indicating a steady level of motoneuronal excitability. Thus, the effects of rTMS are likely to occur at a premotoneuronal site-either at M1 and/or the CM synapse. As the early facilitation reflects activation of direct CM projections, the most likely site of action is the synapse of the CM neurons onto spinal motoneurons.