Local vibration induces changes in spinal and corticospinal excitability in vibrated and antagonist muscles.

Local vibration (LV) applied over the muscle tendon constitutes a powerful stimulus to activate the muscle spindle primary (Ia) afferents that project to the spinal level and are conveyed to the cortical level. This study aimed to identify the neuromuscular changes induced by a 30-min LV-inducing illusions of hand extension on the vibrated flexor carpi radialis (FCR) and the antagonist extensor carpi radialis (ECR) muscles. We studied the change of the maximal voluntary isometric contraction (MVIC, experiment 1) for carpal flexion and extension, motor-evoked potentials (MEPs, experiment 2), cervicomedullary motor-evoked potentials (CMEPs, experiment 2), and Hoffmann's reflex (H-reflex, experiment 3) for both muscles at rest. Measurements were performed before (PRE) and at 0, 30, and 60 min after LV protocol. A lasting decrease in strength was only observed for the vibrated muscle. The reduction in CMEPs observed for both muscles seems to support a decrease in alpha motoneurons excitability. In contrast, a slight decrease in MEPs responses was observed only for the vibrated muscle. The MEP/CMEP ratio increase suggested greater cortical excitability after LV for both muscles. In addition, the H-reflex largely decreased for the vibrated and the antagonist muscles. The decrease in the H/CMEP ratio for the vibrated muscle supported both pre- and postsynaptic causes of the decrease in the H-reflex. Finally, LV-inducing illusions of movement reduced alpha motoneurons excitability for both muscles with a concomitant increase in cortical excitability.NEW & NOTEWORTHY Spinal disturbances confound the interpretation of excitability changes in motor areas and compromise the conclusions reached by previous studies using only a corticospinal marker for both vibrated and antagonist muscles. The time course recovery suggests that the H-reflex perturbations for the vibrated muscle do not only depend on changes in alpha motoneurons excitability. Local vibration induces neuromuscular changes in both vibrated and antagonist muscles at the spinal and cortical levels.

Time of day and sleep effects on motor acquisition and consolidation.

We investigated the influence of the time-of-day and sleep on skill acquisition (i.e., skill improvement immediately after a training-session) and consolidation (i.e., skill retention after a time interval including sleep). Three groups were trained at 10 a.m. (G10am), 3 p.m. (G3pm), or 8 p.m. (G8pm) on a finger-tapping task. We recorded the skill (i.e., the ratio between movement duration and accuracy) before and immediately after the training to evaluate acquisition, and after 24 h to measure consolidation. We did not observe any difference in acquisition according to the time of the day. Interestingly, we found a performance improvement 24 h after the evening training (G8pm), while the morning (G10am) and the afternoon (G3pm) groups deteriorated and stabilized their performance, respectively. Furthermore, two control experiments (G8awake and G8sleep) supported the idea that a night of sleep contributes to the skill consolidation of the evening group. These results show a consolidation when the training is carried out in the evening, close to sleep, and forgetting when the training is carried out in the morning, away from sleep. This finding may have an important impact on the planning of training programs in sports, clinical, or experimental domains.

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.

Too much information is no information: how machine learning and feature selection could help in understanding the motor control of pointing.

The aim of this study was to develop the use of Machine Learning techniques as a means of multivariate analysis in studies of motor control. These studies generate a huge amount of data, the analysis of which continues to be largely univariate. We propose the use of machine learning classification and feature selection as a means of uncovering feature combinations that are altered between conditions. High dimensional electromyogram (EMG) vectors were generated as several arm and trunk muscles were recorded while subjects pointed at various angles above and below the gravity neutral horizontal plane. We used Linear Discriminant Analysis (LDA) to carry out binary classifications between the EMG vectors for pointing at a particular angle, vs. pointing at the gravity neutral direction. Classification success provided a composite index of muscular adjustments for various task constraints-in this case, pointing angles. In order to find the combination of features that were significantly altered between task conditions, we conducted a post classification feature selection i.e., investigated which combination of features had allowed for the classification. Feature selection was done by comparing the representations of each category created by LDA for the classification. In other words computing the difference between the representations of each class. We propose that this approach will help with comparing high dimensional EMG patterns in two ways; (i) quantifying the effects of the entire pattern rather than using single arbitrarily defined variables and (ii) identifying the parts of the patterns that convey the most information regarding the investigated effects.

Muscle effort is best minimized by the right-dominant arm in the gravity field.

The central nervous system (CNS) develops motor strategies that minimize various hidden criteria, such as end-point variance or effort. A large body of literature suggests that the dominant (D) arm is specialized for such open-loop optimization-like processes, whereas the nondominant (ND) arm is specialized for closed-loop postural control. Building on recent results suggesting that the brain plans arm movements that take advantage of gravity effects to minimize muscle effort, the present study tests the hypothesized superiority of the dominant arm motor system for effort minimization. Thirty participants (22.5 ± 2.1 yr old; all right-handed) performed vertical arm movements between two targets (40° amplitude), in two directions (upward and downward) with their two arms (dominant and nondominant). We recorded the arm kinematics and electromyographic activities of the anterior and posterior deltoid to compare two motor signatures of the gravity-related optimization process; i.e., directional asymmetries and negative epochs on phasic muscular activity. We found that these motor signatures were still present during movements performed with the nondominant arm, indicating that the effort-minimization process also occurs for the nondominant motor system. However, these markers were reduced compared with movements performed with the dominant arm. This difference was especially prominent during downward movements, where the optimization of gravity effects occurs early in the movement. Assuming that the dominant arm is optimal to minimize muscle effort, as demonstrated by previous studies, the present results support the hypothesized superiority of the dominant arm motor system for effort minimization.NEW & NOTEWORTHY The hypothesis of hemispheric specialization for feedforward and feedback control has been developed and is still debated. Here, taking advantage of recent knowledge on optimal planning and control of arm movements in the gravity field, we recorded kinematics and electromyographic activities during vertical arm movements performed with the dominant and nondominant arms. Our results reveal that the nondominant arm takes advantage of gravity effects to minimize muscle effort, but less than the dominant arm.

Pain, No Gain: Acute Pain Interrupts Motor Imagery Processes and Affects Mental Training-Induced Plasticity.

Pain influences both motor behavior and neuroplastic adaptations induced by physical training. Motor imagery (MI) is a promising method to recover motor functions, for instance in clinical populations with limited endurance or concomitant pain. However, the influence of pain on the MI processes is not well established. This study investigated whether acute experimental pain could modulate corticospinal excitability assessed at rest and during MI (Exp. 1) and limit the use-dependent plasticity induced by MI practice (Exp. 2). Participants imagined thumb movements without pain or with painful electrical stimulations applied either on digit V or over the knee. We used transcranial magnetic stimulation to measure corticospinal excitability at rest and during MI (Exp. 1) and to evoke involuntary thumb movements before and after MI practice (Exp. 2). Regardless of its location, pain prevented the increase of corticospinal excitability that is classically observed during MI. In addition, pain blocked use-dependent plasticity following MI practice, as testified by a lack of significant posttraining deviations. These findings suggest that pain interferes with MI processes, preventing the corticospinal excitability facilitation needed to induce use-dependent plasticity. Pain should be carefully considered for rehabilitation programs using MI to restore motor function.

Smoothness Discriminates Physical from Motor Imagery Practice of Arm Reaching Movements.

Physical practice (PP) and motor imagery practice (MP) lead to the execution of fast and accurate arm movements. However, there is currently no information about the influence of MP on movement smoothness, nor about which performance parameters best discriminate these practices. In the current study, we assessed motor performances with an arm pointing task with constrained precision before and after PP (n = 15), MP (n = 15), or no practice (n = 15). We analyzed gains between Pre- and Post-Test for five performance parameters: movement duration, mean and maximal velocities, total displacements, and the number of velocity peaks characterizing movement smoothness. The results showed an improvement of performance after PP and MP for all parameters, except for total displacements. The gains for movement duration, and mean and maximal velocities were statistically higher after PP and MP than after no practice, and comparable between practices. However, motor gains for the number of velocity peaks were higher after PP than MP, suggesting that movements were smoother after PP than after MP. A discriminant analysis also identified the number of velocity peaks as the most relevant parameter that differentiated PP from MP. The current results provide evidence that PP and MP specifically modulate movement smoothness during arm reaching tasks. This difference may rely on online corrections through sensory feedback integration, available during PP but not during MP.

Perception of effort and the allocation of physical resources: A generalization to upper-limb motor tasks.

Purpose: The perception of effort (PE) is widely used to prescribe and monitor exercise during locomotor and resistance tasks. The present study examines the validity of PE to prescribe and monitor exercise during upper-limb motor tasks under various loads and speed requirements. Methods: Forty participants volunteered in two experiments. In experiment 1, we used four PE intensities to prescribe exercise on a modified version of the box and block test (BBT) and a pointing task. We investigated the possibility of monitoring the exercise intensity by tracking changes in PE rating in response to three different tempos or additional weights. Experiment 2 replicated the possibility of prescribing the exercise with the PE intensity during the BBT and explored the impact of additional weights on performance and PE during the standardized version of the BBT. Muscle activation, heart rate, and respiratory frequencies were recorded. Results: In experiment 1, increasing the PE intensity to prescribe exercise induced an increased performance between each intensity. Increasing task difficulty with faster movement tempo and adding weight on the forearm increased the rating of PE. Experiment 2 replicated the possibility to use PE intensity for exercise prescription during the BBT. When completing the BBT with an additional weight on the forearm, participants maintained performance at the cost of a higher PE. In both experiments, changes in PE were associated with changes in muscle activation. Conclusion: Our results suggest that PE is a valid tool to prescribe and monitor exercise during upper-limb motor tasks.

Deterioration, Compensation and Motor Control Processes in Healthy Aging, Mild Cognitive Impairment and Alzheimer's Disease.

Aging is associated with modifications of several brain structures and functions. These modifications then manifest as modified behaviors. It has been proposed that some brain function modifications may compensate for some other deteriorated ones, thus maintaining behavioral performance. Through the concept of compensation versus deterioration, this article reviews the literature on motor function in healthy and pathological aging. We first highlight mechanistic studies that used paradigms, allowing us to identify precise compensation mechanisms in healthy aging. Subsequently, we review studies investigating motor function in two often-associated neurological conditions, i.e., mild cognitive impairment and Alzheimer's disease. We point out the need to expand the knowledge gained from descriptive studies with studies targeting specific motor control processes. Teasing apart deteriorated versus compensating processes represents precious knowledge that could significantly improve the prevention and rehabilitation of age-related loss of mobility.

A cross-species neural integration of gravity for motor optimization.

Recent kinematic results, combined with model simulations, have provided support for the hypothesis that the human brain shapes motor patterns that use gravity effects to minimize muscle effort. Because many different muscular activation patterns can give rise to the same trajectory, here, we specifically investigate gravity-related movement properties by analyzing muscular activation patterns during single-degree-of-freedom arm movements in various directions. Using a well-known decomposition method of tonic and phasic electromyographic activities, we demonstrate that phasic electromyograms (EMGs) present systematic negative phases. This negativity reveals the optimal motor plan's neural signature, where the motor system harvests the mechanical effects of gravity to accelerate downward and decelerate upward movements, thereby saving muscle effort. We compare experimental findings in humans to monkeys, generalizing the Effort-optimization strategy across species.

Movement detection thresholds reveal proprioceptive impairments in developmental dyslexia.

Developmental dyslexia is associated with vision and hearing impairments. Whether these impairments are causes or comorbidities is controversial. Because both senses are heavily involved in reading, cognitive theories argue that sensory impairments are comorbidities that result from a lack of reading practice. Sensory theories instead argue that this is sensory impairments that cause reading disabilities. Here we test a discriminant prediction: whether sensory impairments in developmental dyslexia are restrained to reading-related senses or encompass other senses. Sensory theories predict that all senses are affected, whereas, according to the lack of reading practice argument, cognitive theories predict that only reading-related senses are affected. Using a robotic ergometer and fully automatized analyses, we tested proprioceptive acuity in seventeen dyslexic children and seventeen age-matched controls on a movement detection task. Compared to controls, dyslexics had higher and more variable detection thresholds. For the weakest proprioceptive stimuli, dyslexics were twice as long and twice as variable as controls. More, proprioceptive acuity strongly correlated with reading abilities, as measured by blind cognitive evaluations. These results unravel a new sensory impairment that cannot be attributed to a lack of reading practice, providing clear support to sensory theories of developmental dyslexia. Protocol registration: This protocol is part of the following registration, ClinicalTrials.gov Identifier: NCT03364010; December 6, 2017.

Motor Planning of Vertical Arm Movements in Healthy Older Adults: Does Effort Minimization Persist With Aging?

Several sensorimotor modifications are known to occur with aging, possibly leading to adverse outcomes such as falls. Recently, some of those modifications have been proposed to emerge from motor planning deteriorations. Motor planning of vertical movements is thought to engage an internal model of gravity to anticipate its mechanical effects on the body-limbs and thus to genuinely produce movements that minimize muscle effort. This is supported, amongst other results, by direction-dependent kinematics where relative durations to peak accelerations and peak velocity are shorter for upward than for downward movements. The present study compares the motor planning of fast and slow vertical arm reaching movements between 18 young (24 ± 3 years old) and 17 older adults (70 ± 5 years old). We found that older participants still exhibit strong directional asymmetries (i.e., differences between upward and downward movements), indicating that optimization processes during motor planning persist with healthy aging. However, the size of these differences was increased in older participants, indicating that gravity-related motor planning changes with age. We discuss this increase as the possible result of an overestimation of gravity torque or increased weight of the effort cost in the optimization process. Overall, these results support the hypothesis that feedforward processes and, more precisely, optimal motor planning, remain active with healthy aging.

An acute session of motor imagery training induces use-dependent plasticity.

Motor imagery, defined as the mental representation of an action without movement-related sensory inputs, is a well-known intervention to improve motor performance. In the current study, we tested whether use-dependent plasticity, a mechanism underlying motor learning, could be induced by an acute session of motor imagery. By means of transcranial magnetic stimulation (TMS) over the left primary motor cortex, we evoked isolated thumb movements in the right hand and assessed corticospinal excitability in the flexor and extensor pollicis brevis muscles. We measured the mean TMS-induced movement direction before and after an acute session of motor imagery practice. In a first experiment, participants of the imagery group were instructed to repeatedly imagine their thumb moving in a direction deviated by 90° from the pre-test movement. This group, but not the control group, deviated the post-training TMS-induced movements toward the training target direction (+44° ± 62° and -1° ± 23°, respectively). Interestingly, the deviation magnitude was driven by the corticospinal excitability increase in the agonist muscle. In a second experiment, we found that post-training TMS-induced movements were proportionally deviated toward the trained direction and returned to baseline 30 minutes after the motor imagery training. These findings suggest that motor imagery induces use-dependent plasticity and, this neural process is accompanied by corticospinal excitability increase in the agonist muscle.

Direction-dependent arm kinematics reveal optimal integration of gravity cues.

The brain has evolved an internal model of gravity to cope with life in the Earth's gravitational environment. How this internal model benefits the implementation of skilled movement has remained unsolved. One prevailing theory has assumed that this internal model is used to compensate for gravity's mechanical effects on the body, such as to maintain invariant motor trajectories. Alternatively, gravity force could be used purposely and efficiently for the planning and execution of voluntary movements, thereby resulting in direction-depending kinematics. Here we experimentally interrogate these two hypotheses by measuring arm kinematics while varying movement direction in normal and zero-G gravity conditions. By comparing experimental results with model predictions, we show that the brain uses the internal model to implement control policies that take advantage of gravity to minimize movement effort.

Initial information prior to movement onset influences kinematics of upward arm pointing movements.

To elaborate a motor plan and perform online control in the gravity field, the brain relies on priors and multisensory integration of information. In particular, afferent and efferent inputs related to the initial state are thought to convey sensorimotor information to plan the upcoming action. Yet it is still unclear to what extent these cues impact motor planning. Here we examined the role of initial information on the planning and execution of arm movements. Participants performed upward arm movements around the shoulder at three speeds and in two arm conditions. In the first condition, the arm was outstretched horizontally and required a significant muscular command to compensate for the gravitational shoulder torque before movement onset. In contrast, in the second condition the arm was passively maintained in the same position with a cushioned support and did not require any muscle contraction before movement execution. We quantified differences in motor performance by comparing shoulder velocity profiles. Previous studies showed that asymmetric velocity profiles reflect an optimal integration of the effects of gravity on upward movements. Consistent with this, we found decreased acceleration durations in both arm conditions. However, early differences in kinematic asymmetries and EMG patterns between the two conditions signaled a change of the motor plan. This different behavior carried on through trials when the arm was at rest before movement onset and may reveal a distinct motor strategy chosen in the context of uncertainty. Altogether, we suggest that the information available online must be complemented by accurate initial information.

Energy-related optimal control accounts for gravitational load: comparing shoulder, elbow, and wrist rotations.

We permanently deal with gravity force. Experimental evidences revealed that moving against gravity strongly differs from moving along the gravity vector. This directional asymmetry has been attributed to an optimal planning process that optimizes gravity force effects to minimize energy. Yet, only few studies have considered the case of vertical movements in the context of optimal control. What kind of cost is better suited to explain kinematic patterns in the vertical plane? Here, we aimed to understand further how the central nervous system (CNS) plans and controls vertical arm movements. Our reasoning was the following: if the CNS optimizes gravity mechanical effects on the moving limbs, kinematic patterns should change according to the direction and the magnitude of the gravity torque being encountered in the motion. Ten subjects carried out single-joint movements, i.e., rotation around the shoulder (whole arm), elbow (forearm), and wrist (hand) joints, in the vertical plane. Joint kinematics were analyzed and compared with various theoretical optimal model predictions (minimum absolute work-jerk, jerk, torque change, and variance). We found both direction-dependent and joint-dependent variations in several kinematic parameters. Notably, directional asymmetries decreased according to a proximodistal gradient. Numerical simulations revealed that our experimental findings could be attributed to an optimal motor planning (minimum absolute work-jerk) that integrates the direction and the magnitude of gravity torque and minimizes the absolute work of forces (energy-related cost) around each joint. Present results support the general idea that the CNS implements optimal solutions according to the dynamic context of the action.

Prism adaptation by mental practice.

The prediction of our actions and their interaction with the external environment is critical for sensorimotor adaptation. For instance, during prism exposure, which deviates laterally our visual field, we progressively correct movement errors by combining sensory feedback with forward model sensory predictions. However, very often we project our actions to the external environment without physically interacting with it (e.g., mental actions). An intriguing question is whether adaptation will occur if we imagine, instead of executing, an arm movement while wearing prisms. Here, we investigated prism adaptation during mental actions. In the first experiment, participants (n = 54) performed arm pointing movements before and after exposure to the optical device. They were equally divided into six groups according to prism exposure: Prisms-Active, Prisms-Imagery, Prisms-Stationary, Prisms-Stationary-Attention, No Conflict-Prisms-Imagery, No Prisms-Imagery. Adaptation, measured by the difference in pointing errors between pre-test and post-test, occurred only in Prisms-Active and Prisms-Imagery conditions. The second experiment confirmed the results of the first experiment and further showed that sensorimotor adaptation was mainly due to proprioceptive realignment in both Prisms-Active (n = 10) and Prisms-Imagery (n = 10) groups. In both experiments adaptation was greater following actual than imagined pointing movements. The present results are the first demonstration of prism adaptation by mental practice under prism exposure and they are discussed in terms of internal forward models and sensorimotor plasticity.

The temporal structure of vertical arm movements.

The present study investigates how the CNS deals with the omnipresent force of gravity during arm motor planning. Previous studies have reported direction-dependent kinematic differences in the vertical plane; notably, acceleration duration was greater during a downward than an upward arm movement. Although the analysis of acceleration and deceleration phases has permitted to explore the integration of gravity force, further investigation is necessary to conclude whether feedforward or feedback control processes are at the origin of this incorporation. We considered that a more detailed analysis of the temporal features of vertical arm movements could provide additional information about gravity force integration into the motor planning. Eight subjects performed single joint vertical arm movements (45° rotation around the shoulder joint) in two opposite directions (upwards and downwards) and at three different speeds (slow, natural and fast). We calculated different parameters of hand acceleration profiles: movement duration (MD), duration to peak acceleration (D PA), duration from peak acceleration to peak velocity (D PA-PV), duration from peak velocity to peak deceleration (D PV-PD), duration from peak deceleration to the movement end (D PD-End), acceleration duration (AD), deceleration duration (DD), peak acceleration (PA), peak velocity (PV), and peak deceleration (PD). While movement durations and amplitudes were similar for upward and downward movements, the temporal structure of acceleration profiles differed between the two directions. More specifically, subjects performed upward movements faster than downward movements; these direction-dependent asymmetries appeared early in the movement (i.e., before PA) and lasted until the moment of PD. Additionally, PA and PV were greater for upward than downward movements. Movement speed also changed the temporal structure of acceleration profiles. The effect of speed and direction on the form of acceleration profiles is consistent with the premise that the CNS optimises motor commands with respect to both gravitational and inertial constraints.