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.

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.

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.

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.

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.

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.

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.

Online and post-trial feedback differentially affect implicit adaptation to a visuomotor rotation.

Multiple motor learning processes can be discriminated in visuomotor rotation paradigms. At least four processes have been proposed: Implicit adaptation updates an internal model based on prediction errors. Model-free reinforcement reinforces actions that achieve task success. Use-dependent learning favors repetition of prior movements, and strategic learning uses explicit knowledge about the task. The current experiment tested whether the processes involved in motor learning differ when visual feedback is altered. Specifically, we hypothesized that online and post-trial feedback would cause different amounts of implicit adaptation. Twenty subjects performed drawing movements to targets under a 45° counterclockwise visuomotor rotation while aiming at a clockwise adjacent target. Subjects received visual feedback via a cursor on a screen. One group saw the cursor throughout the movement (online feedback), while the other only saw the final position after movement execution (post-trial feedback). Both groups initially hit the target by applying the strategy. After 80 trials, subjects with online feedback had drifted in clockwise direction [mean direction error: 15.1° (SD 11.2°)], thus overcompensating the rotation. Subjects with post-trial feedback remained accurate [mean: 0.7° (SD 2.0°), TIME × GROUP: F = 3.926, p = 0.003]. We interpret this overcompensation to reflect implicit adaptation isolated from other mechanisms, because it is driven by prediction error rather than task success (model-free reinforcement) or repetition (use-dependent learning). The current findings extend previous work (e.g., Mazzoni and Krakauer in J Neurosci 26:3642-3645, 2006; Hinder et al. in Exp Brain Res 201:191-207, 2010) and suggest that online feedback promotes more implicit adaptation than does post-trial feedback.

Specific interpretation of augmented feedback changes motor performance and cortical processing.

It is well established that the presence of external feedback, also termed augmented feedback, can be used to improve performance of a motor task. The present study aimed to elucidate whether differential interpretation of the external feedback signal influences the time to task failure of a sustained submaximal contraction and modulates motor cortical activity. In Experiment 1, subjects had to maintain a submaximal contraction (30% of maximum force) performed with their thumb and index finger. Half of the tested subjects were always provided with feedback about joint position (pF-group), whereas the other half of the subjects were always provided with feedback about force (fF-group). Subjects in the pF-group were led to belief in half of their trials that they would receive feedback about the applied force, and subjects in the fF-group to receive feedback about the position. In both groups (fF and pF), the time to task failure was increased when subjects thought to receive feedback about the force. In Experiment 2, subthreshold transcranial magnetic stimulation was applied over the right motor cortex and revealed an increased motor cortical activity when subjects thought to receive feedback about the joint position. The results showed that the interpretation of feedback influences motor behavior and alters motor cortical activity. The current results support previous studies suggesting a distinct neural control of force and position.

Cutaneous mechanisms of isometric ankle force control.

The sense of force is critical in the control of movement and posture. Multiple factors influence our perception of exerted force, including inputs from cutaneous afferents, muscle afferents and central commands. Here, we studied the influence of cutaneous feedback on the control of ankle force output. We used repetitive electrical stimulation of the superficial peroneal (foot dorsum) and medial plantar nerves (foot sole) to disrupt cutaneous afferent input in 8 healthy subjects. We measured the effects of repetitive nerve stimulation on (1) tactile thresholds, (2) performance in an ankle force-matching and (3) an ankle position-matching task. Additional force-matching experiments were done to compare the effects of transient versus continuous stimulation in 6 subjects and to determine the effects of foot anesthesia using lidocaine in another 6 subjects. The results showed that stimulation decreased cutaneous sensory function as evidenced by increased touch threshold. Absolute dorsiflexion force error increased without visual feedback during peroneal nerve stimulation. This was not a general effect of stimulation because force error did not increase during plantar nerve stimulation. The effects of transient stimulation on force error were greater when compared to continuous stimulation and lidocaine injection. Position-matching performance was unaffected by peroneal nerve or plantar nerve stimulation. Our results show that cutaneous feedback plays a role in the control of force output at the ankle joint. Understanding how the nervous system normally uses cutaneous feedback in motor control will help us identify which functional aspects are impaired in aging and neurological diseases.

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.

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.

Pathway-specific plasticity in the human spinal cord.

The aim of the present study was to artificially induce plasticity in the human spinal cord and evaluate whether this plasticity is pathway specific. For this purpose, a technique called paired associative stimulation (PAS) was applied. Volleys evoked by transcranial magnetic stimulation over the primary motor cortex and peripheral nerve stimulation of the nervus tibialis in the popliteal fossa were timed to coincide at the spinal level. The transmission of different corticospinal projections was assessed before and after PAS using conditioned H-reflexes. Different groups of healthy volunteers (28 ± 5 years) were tested; intervention groups 1 (n = 9) and 2 (n = 8) received spinal PAS (360 paired stimuli) and the induced effects were evaluated using cortical (group 1) or cervicomedullary (group 2) conditioning of musculus soleus H-reflexes. After spinal PAS, the conditioned H-reflexes were significantly facilitated when tested with cortical and cervicomedullary stimulation. The effect of the latter technique is independent of changes in the excitability of cortical neurons. Therefore, the finding that conditioned H-reflexes were increased after spinal PAS when tested with both cortical and cervicomedullary stimulation suggests that neural plasticity was induced within the spinal cord. The facilitation could only be observed for specific inter-stimulus intervals between volleys induced by peripheral nerve stimulation and transcranial magnetic stimulation. As the specific inter-stimulus intervals were assumed to relate to transmission within specific motor pathways, it is argued that changes in the corticospinal transmission were pathway-specific. These findings may be helpful in inducing and assessing neural plasticity in pathological conditions like spinal cord injuries.

How neurons make us jump: the neural control of stretch-shortening cycle movements.

How can the human central nervous system (CNS) control complex jumping movements task- and context-specifically? This review highlights the complex interaction of multiple hierarchical levels of the CNS, which work together to enable stretch-shortening cycle contractions composed of activity resulting from feedforward (preprogrammed) and feedback (reflex) loops.

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.

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.

Contribution of afferent feedback and descending drive to human hopping.

During hopping an early burst can be observed in the EMG from the soleus muscle starting about 45 ms after touch-down. It may be speculated that this early EMG burst is a stretch reflex response superimposed on activity from a supra-spinal origin. We hypothesised that if a stretch reflex indeed contributes to the early EMG burst, then advancing or delaying the touch-down without the subject's knowledge should similarly advance or delay the burst. This was indeed the case when touch-down was advanced or delayed by shifting the height of a programmable platform up or down between two hops and this resulted in a correspondent shift of the early EMG burst. Our second hypothesis was that the motor cortex contributes to the first EMG burst during hopping. If so, inhibition of the motor cortex would reduce the magnitude of the burst. By applying a low-intensity magnetic stimulus it was possible to inhibit the motor cortex and this resulted in a suppression of the early EMG burst. These results suggest that sensory feedback and descending drive from the motor cortex are integrated to drive the motor neuron pool during the early EMG burst in hopping. Thus, simple reflexes work in concert with higher order structures to produce this repetitive movement.

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.