Gravity-efficient motor control is associated with contraction-dependent intracortical inhibition.

In humans, moving efficiently along the gravity axis requires shifts in muscular contraction modes. Raising the arm up involves shortening contractions of arm flexors, whereas the reverse movement can rely on lengthening contractions with the help of gravity. Although this control mode is universal, the neuromuscular mechanisms that drive gravity-oriented movements remain unknown. Here, we designed neurophysiological experiments that aimed to track the modulations of cortical, spinal, and muscular outputs of arm flexors during vertical movements with specific kinematics (i.e., optimal motor commands). We report a specific drop of corticospinal excitability during lengthening versus shortening contractions, with an increase of intracortical inhibition and no change in spinal motoneuron responsiveness. We discuss these contraction-dependent modulations of the supraspinal motor output in the light of feedforward mechanisms that may support gravity-tuned motor control. Generally, these results shed a new perspective on the neural policy that optimizes movement control along the gravity axis.

Children with Dyslexia Have Altered Cross-Modal Processing Linked to Binocular Fusion. A Pilot Study.

INTRODUCTION: The cause of dyslexia, a reading disability characterized by difficulties with accurate and/or fluent word recognition and by poor spelling and decoding abilities, is unknown. A considerable body of evidence shows that dyslexics have phonological disorders. Other studies support a theory of altered cross-modal processing with the existence of a pan-sensory temporal processing deficit associated with dyslexia. Learning to read ultimately relies on the formation of automatic multisensory representations of sounds and their written representation while eyes fix a word or move along a text. We therefore studied the effect of brief sounds on vision with a modification of binocular fusion at the same time (using the Maddox Rod test). METHODS: To check if the effect of sound on vision is specific, we first tested with sounds and then replaced them with proprioceptive stimulation on 8 muscular sites. We tested two groups of children composed respectively of 14 dyslexic children and 10 controls. RESULTS: The results show transient visual scotoma (VS) produced by sensory stimulations associated with the manipulation of oculomotor balance, the effect being drastically higher in the dyslexic group. The spatial distribution of the VS is stochastic. The effect is not specific for sounds but exists also with proprioceptive stimulations. DISCUSSION: Although there was a very significant difference between the two groups, we were not able to correlate the (VS) occurrence with the dyslexic's reading performance. One possibility to confirm the link between VS and reading impairment would be to find a specific treatment reducing the occurrence of the VS and to check its effect on dyslexia.

Presynaptic inhibition mechanisms may subserve the spinal excitability modulation induced by neuromuscular electrical stimulation.

This study aimed at deciphering the origins of spinal excitability modulation that follows neuromuscular electrical stimulation (NMES). Ten participants (age: 24.6 ±â€¯4.2 years) performed 2 randomized NMES sessions on plantar flexors with frequencies of stimulations of 20 or 100 Hz (pulse width: 1 ms) at 20% of maximal voluntary contraction (MVC). Before and after each session, the posterior tibial nerve was stimulated to record H-reflex of soleus (SOL), gastrocnemius medialis (GM) and gastrocnemius lateralis (GL). D1 presynaptic inhibition was assessed by conditioning H reflex with prior common peroneal nerve stimulation. Resting H-reflex of SOL decreased after both protocols, but in a greater extent following the 100 Hz session (100 Hz: -34.6 ±â€¯7.3%, 20 Hz: -17.1 ±â€¯3.8%; P = 0.002), accompanied by an increase of presynaptic inhibition (+22 ±â€¯5.8% at 100 Hz vs. +8 ± 3.7% at 20 Hz, P < 0.001). GM and GL spinal excitability and presynaptic inhibition were also altered after NMES, but in a similarly extent after 20 Hz and 100 Hz protocols. Neuromuscular fatigue following a single session of NMES involves spinal presynaptic circuitry, even at low stimulation frequency. The spinal sensitivity to NMES seems also muscle dependent.

Daily modulation of the speed-accuracy trade-off.

Goal-oriented arm movements are characterized by a balance between speed and accuracy. The relation between speed and accuracy has been formalized by Fitts' law and predicts a linear increase in movement duration with task constraints. Up to now this relation has been investigated on a short-time scale only, that is during a single experimental session, although chronobiological studies report that the motor system is shaped by circadian rhythms. Here, we examine whether the speed-accuracy trade-off could vary during the day. Healthy adults carried out arm-pointing movements as accurately and fast as possible toward targets of different sizes at various hours of the day, and variations in Fitts' law parameters were scrutinized. To investigate whether the potential modulation of the speed-accuracy trade-off has peripheral and/or central origins, a motor imagery paradigm was used as well. Results indicated a daily (circadian-like) variation for the durations of both executed and mentally simulated movements, in strictly controlled accuracy conditions. While Fitts' law was held for the whole sessions of the day, the slope of the relation between movement duration and task difficulty expressed a clear modulation, with the lowest values in the afternoon. This variation of the speed-accuracy trade-off in executed and mental movements suggests that, beyond execution parameters, motor planning mechanisms are modulated during the day. Daily update of forward models is discussed as a potential mechanism.

Central Contribution to Electrically Induced Fatigue depends on Stimulation Frequency.

PURPOSE: This study analyzed the impact of several protocols of neuromuscular electrical stimulation (NMES), matched with a similar total torque-time integral, on muscle activation pathways and neuromuscular fatigue. METHODS: Ten young healthy participants (age, 24.6 ± 4.2 yr) performed three randomized NMES sessions on the triceps surae muscles with 20-, 60-, or 100-Hz stimulation frequencies (pulse duration, 1 ms), with pulse amplitude (IES) set at 20% of isometric maximal voluntary contraction (MVC). Muscle activity during NMES was assessed by means of the twitch, the soleus H-reflex and M wave responses evoked by single muscle stimulation at IES. Neuromuscular fatigue was assessed as the changes in evoked and MVC torques and the underlying mechanisms by analyzing variations in superimposed maximal M-waves (Msup), normalized H-reflexes (Hsup/Msup) and V-waves (V/Msup) of the triceps surae muscles. RESULTS: Electromyographic responses at IES suggested that the relative contribution of the indirect muscle activation increases as the stimulation frequency was high and the pulse amplitude was low (P = 0.03). The decrease in MVC torque after NMES was significantly (P = 0.003) greater after 100-Hz protocol (20 Hz, -9.6% ± 3.3%; 60 Hz, -10.7% ± 3.2%; 100 Hz, -16.3% ± 2.7%). Hsup/Msup decreased significantly (P < 0.01) by 31% ± 4% after the 100-Hz protocol only and V/Msup decreased significantly (P <0.05) after both 60- and 100-Hz protocols. CONCLUSIONS: The combination of high-stimulation frequencies and low-pulse amplitude induced the greatest neuromuscular fatigue. Low frequencies (20 Hz) induced alterations mainly at the muscle level, whereas higher frequencies (60 to 100 Hz) rather induced modulations at both spinal and supraspinal levels.

High-frequency neuromuscular electrical stimulation modulates interhemispheric inhibition in healthy humans.

High-frequency neuromuscular electrical stimulation (HF NMES) induces muscular contractions through neural mechanisms that partially match physiological motor control. Indeed, a portion of the contraction arises from central mechanisms, whereby spinal motoneurons are recruited through the evoked sensory volley. However, the involvement of supraspinal centers of motor control during such stimulation remains poorly understood. Therefore, we tested whether a single HF NMES session applied to the upper limb influences interhemispheric inhibition (IHI) from left to right motor cortex (M1). Using noninvasive electrophysiology and transcranial magnetic stimulation, we evaluated the effects of a 10-min HF NMES session applied to a right wrist flexor on spinal and corticospinal excitability of both arms, as well as IHI, in healthy subjects. HF NMES induced a rapid decline in spinal excitability on the right stimulated side that closely matched the modulation of evoked force during the protocol. More importantly, IHI was significantly increased by HF NMES, and this increase was correlated to the electromyographic activity within the contralateral homologous muscle. Our study highlights a new neurophysiological mechanism, suggesting that HF NMES has an effect on the excitability of the transcallosal pathway probably to regulate the lateralization of the motor output. The data suggest that HF NMES can modify the hemispheric balance between both M1 areas. These findings provide important novel perspectives for the implementation of HF NMES in sport training and neurorehabilitation. NEW & NOTEWORTHY: High-frequency neuromuscular electrical stimulation (HF NMES) induces muscular contractions that partially match physiological motor control. Here, we tested whether HF NMES applied to the upper limb influences interhemispheric inhibition. Our results show that interhemispheric inhibition was increased after HF NMES and that this increase was correlated to the electromyographic activity within the contralateral homologous muscle. This opens up original perspectives for the implementation of HF NMES in sport training and neurorehabilitation.

Interhemispheric inhibition is dynamically regulated during action observation.

It is now well established that the motor system plays a pivotal role in action observation and that the neurophysiological processes underlying perception and action overlaps. However, while various experiments have shown a specific facilitation of the contralateral motor cortex during action observation, no information is available concerning the dynamics of interhemispheric interactions. The aim of the present study was, therefore, to assess interhemispheric inhibition during the observation of others' actions. We designed a transcranial magnetic stimulation (TMS) experiment in which we measured both corticospinal excitability and interhemispheric inhibition, this latter by means of the ipsilateral silent period (iSP), while participants observed two motor tasks (tapping or grasping). We show that the iSP is enhanced during movement observation and that this modulation is tuned to the kinematics of the observed movements. An additional experiment was performed in which the TMS intensity was adjusted to match corticospinal excitability between rest and action observation. This resulted in a relative decrease of iSP. Overall, our data strongly suggest that action observation, as action execution, involves interhemispheric inhibitory mechanisms between the two motor cortices, and that this neural activity appears to be firmly shaped by the ongoing observed movement and its intrinsic dynamics.

Daily update of motor predictions by physical activity.

Motor prediction, i.e., the ability to predict the sensory consequences of motor commands, is critical for adapted motor behavior. Like speed or force, the accuracy of motor prediction varies in a 24-hour basis. Although the prevailing view is that basic biological markers regulate this circadian modulation, behavioral factors such as physical activity, itself modulated by the alternation of night and day, can also regulate motor prediction. Here, we propose that physical activity updates motor prediction on a daily basis. We tested our hypothesis by up- and down-regulating physical activity via arm-immobilization and high-intensity training, respectively. Motor prediction was assessed by measuring the timing differences between actual and mental arm movements. Results show that although mental movement time was modulated during the day when the arm was unconstrained, it remained constant when the arm was immobilized. Additionally, increase of physical activity, via release from immobilization or intense bout of training, significantly reduced mental movement time. Finally, mental and actual times were similar in the afternoon in the unconstrained condition, indicating that predicted and actual movements match after sufficient amount of physical activity. Our study supports the view that physical activity calibrates motor predictions on a daily basis.

Motor cortical plasticity induced by motor learning through mental practice.

Several investigations suggest that actual and mental actions trigger similar neural substrates. Motor learning via physical practice results in long-term potentiation (LTP)-like plasticity processes, namely potentiation of M1 and a temporary occlusion of additional LTP-like plasticity. However, whether this neuroplasticity process contributes to improve motor performance through mental practice remains to be determined. Here, we tested skill learning-dependent changes in primary motor cortex (M1) excitability and plasticity by means of transcranial magnetic stimulation (TMS) in subjects trained to physically execute or mentally perform a sequence of finger opposition movements. Before and after physical practice and motor-imagery practice, M1 excitability was evaluated by measuring the input-output (IO) curve of motor evoked potentials. M1 LTP and long-term depression (LTD)-like plasticity was assessed with paired-associative stimulation (PAS) of the median nerve and motor cortex using an interstimulus interval of 25 ms (PAS25) or 10 ms (PAS10), respectively. We found that even if after both practice sessions subjects significantly improved their movement speed, M1 excitability and plasticity were differentially influenced by the two practice sessions. First, we observed an increase in the slope of IO curve after physical but not after MI practice. Second, there was a reversal of the PAS25 effect from LTP-like plasticity to LTD-like plasticity following physical and MI practice. Third, LTD-like plasticity (PAS10 protocol) increased after physical practice, whilst it was occluded after MI practice. In conclusion, we demonstrated that MI practice lead to the development of neuroplasticity, as it affected the PAS25- and PAS10- induced plasticity in M1. These results, expanding the current knowledge on how MI training shapes M1 plasticity, might have a potential impact in rehabilitation.

Direct mapping rather than motor prediction subserves modulation of corticospinal excitability during observation of actions in real time.

Motor facilitation refers to the specific increment in corticospinal excitability (CSE) elicited by the observation of actions performed by others. To date, the precise nature of the mechanism at the basis of this phenomenon is unknown. One possibility is that motor facilitation is driven by a predictive process reminiscent of the role of forward models in motor control. Alternatively, motor facilitation may result from a model-free mechanism by which the basic elements of the observed action are directly mapped onto their cortical representations. Our study was designed to discern these alternatives. To this aim, we recorded the time course of CSE for the first dorsal interosseous (FDI) and the abductor digiti minimi (ADM) during observation of three grasping actions in real time, two of which strongly diverged in kinematics from their natural (invariant) form. Although artificially slow movements used in most action observation studies might enhance the observer's discrimination performance, the use of videos in real time is crucial to maintain the time course of CSE within the physiological range of daily actions. CSE was measured at 4 time points within a 240-ms window that best captured the kinematic divergence from the invariant form. Our results show that CSE of the FDI, not the ADM, closely follows the functional role of the muscle despite the mismatch between the natural and the divergent kinematics. We propose that motor facilitation during observation of actions performed in real time reflects the model-free coding of perceived movement following a direct mapping mechanism.

Observing and perceiving: A combined approach to induce plasticity in human motor cortex.

OBJECTIVE: To test whether action observation combined with peripheral nerve electrical stimulation was able to evoke plasticity in the primary motor cortex (M1). METHODS: The stimulation protocol consisted in the observation of a video showing repetitive thumb-index tapping movements (AO) combined with peripheral electrical nerve stimulation (PNS) delivered on the median nerve (AO-PNS). M1 excitability, measured by means of transcranial magnetic stimulation, was compared with that assessed after AO and PNS alone. RESULTS: M1 excitability increased after AO-PNS, whilst no modifications occurred after AO and PNS alone. The increased M1 excitability after AO-PNS was long-lasting (45 min) and specific for the stimulated muscle. CONCLUSIONS: This study described an innovative stimulation paradigm that exploited the mirror neuron system to induce plasticity in M1. However, this occurred only when action observation was combined with afferent signals coming from periphery. SIGNIFICANCE: This study supports the literature proposing the mirror neuron system as neural substrate for rehabilitation and opens a debate on the rehabilitative treatments that employ AO to improve patients' motor functions. Indeed, these results suggest that AO has to be combined with afferent inputs from periphery to evoke plasticity in the human motor system.

Dominant vs. nondominant arm advantage in mentally simulated actions in right handers.

Although plentiful data are available regarding mental states involving the dominant-right arm, the evidence for the nondominant-left arm is sparse. Here, we investigated whether right-handers can generate accurate predictions with either the right or the left arm. Fifteen adults carried out actual and mental arm movements in two directions with varying inertial resistance (inertial anisotropy phenomenon). We recorded actual and mental movement times and used the degree of their similarity as an indicator for the accuracy of motor imagery/prediction process. We found timing correspondences (isochrony) between actual and mental right arm movements in both rightward (low inertia resistance) and leftward (high inertia resistance) directions. Timing similarities between actual and mental left arm movements existed for the leftward direction (low inertia resistance) but not for the rightward direction (high inertia resistance). We found similar results when participants reaching towards the midline of the workspace, a result that excludes a hemispace effect. Electromyographic analysis during mental movements showed that arm muscles remained inactivate, thus eliminating a muscle activation strategy that could explain intermanual differences. Furthermore, motor-evoked potentials enhancement in both right and left biceps brachii during mental actions indicated that subjects were actively engaged in mental movement simulation and that the disadvantage of the left arm cannot be attributed to the nonactivation of the right motor cortex. Our findings suggest that predictive mechanisms are more robust for the right than the left arm in right-handers. We discussed these findings from the perspective of the internal models theory and the dynamic-dominance hypothesis of laterality.

Interhemispheric inhibition during mental actions of different complexity.

Several investigations suggest that actual and mental actions trigger similar neural substrates. Yet, neurophysiological evidences on the nature of interhemispheric interactions during mental movements are still meagre. Here, we asked whether the content of mental images, investigated by task complexity, is finely represented in the inhibitory interactions between the two primary motor cortices (M1s). Subjects' left M1 was stimulated by means of transcranial magnetic stimulation (TMS) while they were performing actual or mental movements of increasing complexity with their right hand and exerting a maximum isometric force with their left thumb and index. Thus, we simultaneously assessed the corticospinal excitability in the right opponent pollicis muscle (OP) and the ipsilateral silent period (iSP) in the left OP during actual and mental movements. Corticospinal excitability in right OP increased during actual and mental movements, but task complexity-dependent changes were only observed during actual movements. Interhemispheric motor inhibition in the left OP was similarly modulated by task complexity in both mental and actual movements. Precisely, the duration and the area of the iSP increased with task complexity in both movement conditions. Our findings suggest that mental and actual movements share similar inhibitory neural circuits between the two homologous primary motor cortex areas.

Time-of-day effects on the internal simulation of motor actions: psychophysical evidence from pointing movements with the dominant and non-dominant arm.

It is well known that circadian rhythms modulate human physiology and behavior at various levels. However, chronobiological data concerning mental and sensorimotor states of motor actions are still lacking in the literature. In the present study, we examined the effects of time-of-day on two important aspects of the human motor behavior: prediction and laterality. Motor prediction was experimentally investigated by means of imagined movements and laterality by comparing the difference in temporal performance between right and left arm movements. Ten healthy participants had to actually perform or to imagine performing arm-pointing movements between two targets at different hours of the day (i.e., 08:00, 11:00, 14:00, 17:00, 20:00, and 23:00 h). Executed and imagined movements were accomplished with both the right and left arm. We found that both imagined and executed arm pointing movements significantly fluctuated through the day. Furthermore, the accuracy of motor prediction, investigated by the temporal discrepancy between executed and imagined movements, was significantly better in the afternoon (i.e., 14:00, 17:00, and 20:00 h) than morning (08:00 and 11:00 h) and evening (23:00 h). Our results also revealed that laterality was not stable throughout the day. Indeed, the smallest temporal differences between the two arms appeared at 08:00 and 23:00 h, whereas the largest ones occurred at the end of the morning (11:00 h). The daily variation of motor imagery may suggest that internal predictive models are flexible entities that are continuously updated throughout the day. Likewise, the variations in temporal performance between the right and the left arm during the day may indicate a relative independence of the two body sides in terms of circadian rhythms. In general, our findings suggest that cognitive (i.e., mental imagery) and motor (i.e., laterality) states of human behavior are modulated by circadian rhythms.

Circadian modulation of mentally simulated motor actions: implications for the potential use of motor imagery in rehabilitation.

BACKGROUND: . Mental practice through motor imagery improves subsequent motor performance and thus mental training is considered to be a potential tool in neuromotor rehabilitation. OBJECTIVE: . The authors investigated whether a circadian fluctuation of the motor imagery process occurs, which could be relevant in scheduling mental training in rehabilitation programs. METHODS: . The executed and imagined durations of walking and writing movements were recorded every 3 hours from 8 AM to 11 PM in healthy participants. The authors made a cosinor analysis on the temporal features of these movements to detect circadian rhythms. Temporal differences between executed and imagined movements as well as their variability during the day were also quantified. RESULTS: . Circadian rhythms were detected for both the executed and the imagined movements. Furthermore, these rhythms covaried between them and with body temperature. The participants' ability to internally simulate their movements also fluctuated significantly during the day. The isochrony between the executed and the imagined movements was exclusively observed between 2 PM and 8 PM. In the morning (8 AM and 11 AM) and the evening (11 PM), the durations of the imagined movements were significantly longer than the durations of executed movements. CONCLUSIONS: . Predictive internal models fluctuate in a circadian basis, as do many other physiological parameters. It could be important to take into consideration the time of day in the planning of rehabilitation programs using physical or mental training.

The influence of eye movements on the temporal features of executed and imagined arm movements.

The very close coordination between eye and hand indicates that eye movements are parts of the neural processes underlying the planning and control of arm movements. Eye movements are fundamental during observed actions and play a functional role in visual mental imagery. However, the role of eye movements during imagined actions is still unknown. Here, we report the timing features of eye and arm pointing movements for nine healthy participants in four conditions: Executed movements with orientation saccades (Eyes Free) or with no saccades (Eyes Motionless), and Imagined movements with Eyes Free or with Eyes Motionless. The first result was a facilitation effect of saccades upon both executed and imagined arm movements as revealed by the shorter arm movement durations in Eyes Free than in Eyes Motionless. Another interesting finding was that executed and imagined movements preserved their temporal similarities in both Eyes Free and Eyes Motionless, suggesting that the accuracy of motor representations is not dependent on the presence or lack of eye movements. This result and the close similarities between the EOG patterns accompanying both executed and imagined arm pointing movements in Eyes Free, argue in favour of a similar neurocognitive network in executed and imagined actions. We propose that internal forward models provide fine estimations of the temporal features of imagined arm movements, whatever they accompanied, or not, by orientation saccades.