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In the present study, we examined the role of the cerebellum in temporal adaptive learning during a coincident timing task, i.e., a baseball-like hitting task involving a moving ball presented on a computer monitor. The subjects were required to change the timing of their responses based on imposed temporal perturbations. Using paired-pulse transcranial magnetic stimulation, we measured cerebellar brain inhibition (CBI) before, during, and after the temporal adaptive learning. Reductions in CBI only occurred during and after the temporal adaptive learning, regardless of the direction of the temporal perturbations. In addition, the changes in CBI were correlated with the magnitude of the adaptation. Here, we showed that the cerebellum is essential for learning about and controlling the timing of movements during temporal adaptation. Furthermore, changes in cerebellar-primary motor cortex connectivity occurred during temporal adaptation, as has been previously reported for spatial adaptation.

Acquisition of new motor skills induces plastic reorganization in the primary motor cortex (M1). Previous studies have demonstrated the increases in the M1 excitability through motor skill learning. However, this M1 reorganization is highly variable between individuals even though they improve their skill performance through the same training protocol. To reveal the source of this interindividual variability, we examined the relationship between an acquisition of memory-guided feedforward movements and the learning-induced increases in the M1 excitability. Twenty-eight subjects participated in experiment 1. We asked subjects to learn a visuomotor tracking task. The subjects controlled a cursor on a PC monitor to pursue a target line by performing ankle dorsiflexion and plantar flexion. In experiment 1, we removed the online visual feedback provided by the cursor movement once every six trials, which enabled us to assess whether the subjects could perform accurate memory-guided movements. Motor-evoked potentials (MEP) were elicited in the tibialis anterior muscle by transcranial magnetic stimulation of the relevant M1 before and after the learning of the visuomotor tracking task and after half the trials. We found that the MEP amplitude was increased along with the improvement in memory-guided movements. In experiment 2 ( n = 10), we confirmed this relationship by examining whether the improvement in memory-guided movements induces increases in MEP amplitude. The results of this study indicate that the plastic reorganization of the M1 induced by the learning of a visuomotor skill is associated with the acquisition of memory-guided movements. NEW & NOTEWORTHY Acquisition of novel motor skills increases excitability of the primary motor cortex (M1). We recently reported that the amount of increases in the M1 excitability is highly variable between individuals even though they learned the same skill to the similar extent, yet the sources of this interindividual variability still remain unclear. The present study revealed that this interindividual variability is associated with whether individuals acquire a motor memory, which enables them to produce accurate memory-guided movements.

Dexterous finger movements are often characterized by highly coordinated movements. Such coordination might be derived from reorganization of the corticospinal system. In this study, we investigated 1) the manner in which finger movement covariation patterns are acquired, by examining the effects of the implicit and explicit learning of a serial reaction time task (SRTT), and 2) how such changes in finger coordination are represented in the corticospinal system. The subjects learned a button press sequence in both implicit and explicit learning conditions. In the implicit conditions, they were naive about what they were learning, whereas in the explicit conditions the subjects consciously learned the order of the sequence elements. Principal component analysis decomposed both the voluntary movements produced during the SRTT and the passive movements evoked by transcranial magnetic stimulation (TMS) over the primary motor cortex into a set of five finger joint covariation patterns. The structures of the voluntary and passive TMS-evoked movement patterns were reorganized by implicit learning but not explicit learning. Furthermore, in the implicit learning conditions the finger covariation patterns derived from the TMS-evoked and voluntary movements spanned similar movement subspaces. These results provide the first evidence that skilled sequential finger movements are acquired differently through implicit and explicit learning, i.e., the changes in finger coordination patterns induced by implicit learning are accompanied by functional reorganization of the corticospinal system, whereas explicit learning results in faster recruitment of individual finger movements without causing any changes in finger coordination. NEW & NOTEWORTHY Skilled sequential multifinger movements are characterized as highly coordinated movement patterns. These finger coordination patterns are represented in the corticospinal system, yet it still remains unclear how these patterns are acquired through implicit and explicit motor sequence learning. A direct comparison of learning-related changes between actively generated finger movements and passively evoked finger movements by TMS provided evidence that finger coordination patterns represented in the corticospinal system are reorganized through implicit, but not explicit, sequence learning.

To examine the factors that influence the inter-individual differences in the changes in primary motor cortex (M1) excitability seen after motor learning, we investigated the relationship between the amplitude of transcranial magnetic stimulation-induced motor evoked potentials (MEP) and short-latency afferent inhibition (SAI) after motor learning, based on the working hypothesis that SAI can be used to evaluate cortical acetylcholine (ACh) activity. To confirm this working hypothesis, we manipulated the arousal state of the subjects using a vigilance task, the outcomes of which might be correlated with cortical ACh activity, and investigated the effects of arousal state on SAI. As a result, we showed that SAI was significantly affected by arousal state. Consequently, we concluded that the subjects’ arousal state during motor learning tasks is one of factors to influence on inter-individual differences in the changes in M1 excitability seen after motor learning tasks.

How dexterous finger movements are acquired by the nervous system is a fundamental question in the neuroscience field. Previous studies have demonstrated that finger movements can be decomposed into finger covariation patterns, and these patterns are represented in the corticospinal system. However, it remains unclear how such covariation patterns represented in the corticospinal system develop during the acquisition of novel finger movements. In this study, each subject learned to perform a novel finger movement, which was mapped to a region outside the movement subspace spanned by common finger movements seen in daily life, through a custom task. After subjects practiced the task, we detected changes in the finger covariation patterns derived from artificially (transcranial magnetic stimulation) evoked finger joint movements. The artificially evoked movement-derived patterns seen after the training period were associated with both the novel and common finger movements. Regarding the patterns extracted from the artificially evoked movements, the number required to explain most of the variance in the data was unchanged after the training period. Our results indicate that novel finger movements are acquired through the reorganization of preexisting finger covariation patterns represented in the corticospinal system rather than the development of new patterns. These findings might have implications for the basic mechanism responsible for the development of movement repertories in the nervous system. NEW & NOTEWORTHY Various types of finger movements involve common finger covariation patterns, and these patterns are represented in the corticospinal system. Here we examined how a novel finger covariation pattern is acquired in that system through training of a novel finger movement that is irrelevant to common finger movements. Using transcranial magnetic stimulation, we found that the preexisting patterns that contribute to finer control of finger movements are rapidly reorganized to encode the novel pattern through the training.

Background Previous studies have shown that primary motor cortex (M1) excitability is modulated by motor skill learning and that the M1 plays a crucial role in motor memory. However, the following questions remain: 1) At what stage do changes in M1 excitability occur? 2) Are learning-induced changes in leg M1 excitability associated with motor memory? Here, we did two experiments to answer these questions. Methods and results In experiment 1, subjects learned a visuomotor tracking task over two consecutive days. Before and after the task in Day 1, we recorded input–output curves of the motor evoked potentials (I–O curve) produced in the tibialis anterior muscle by transcranial magnetic stimulation. We found that the changes in M1 excitability were affected by learning stage. In addition, the changes in M1 excitability in Day 1 were correlated with the retention. In experiment 2, we recorded I-O curves before learning, after the fast-learning stage, and after learning. We found no changes in M1 excitability immediately after the fast-learning stage. Furthermore, a significant relationship between the length of slow-learning stage and the changes in M1 excitability was detected. Conclusions Previous studies have suggested that optimal motor commands are repeatedly used during the slow-learning stage. Therefore, present results indicate that changes in M1 excitability occur during the slow-learning stage and that such changes are proportional to motor skill retention because use-dependent plasticity occur by repetitive use of same motor commands during the slow-learning stage.

Patterned sensory nerve stimulation has been shown to induce plastic changes in the reciprocal Ia inhibitory circuit. However, the mechanisms underlying these changes have not yet been elucidated in detail. The aim of the present study was to determine whether the reactivity of Ia inhibitory interneurons could be altered by patterned sensory nerve stimulation. The degree of reciprocal Ia inhibition, the conditioning effects of transcranial magnetic stimulation (TMS) on the soleus (SOL) muscle H-reflex, and the ratio of the maximum H-reflex amplitude versus maximum M-wave (H(max)/M(max)) were examined in 10 healthy individuals. Patterned electrical nerve stimulation was applied to the common peroneal nerve every 1 s (100 Hz-5 train) at the motor threshold intensity of tibialis anterior muscle to induce activity changes in the reciprocal Ia inhibitory circuit. Reciprocal Ia inhibition, the TMS-conditioned H-reflex amplitude, and H(max)/M(max) were recorded before, immediately after, and 15 min after the electrical stimulation. The patterned electrical nerve stimulation significantly increased the degree of reciprocal Ia inhibition and decreased the amplitude of the TMS-conditioned H-reflex in the short-latency inhibition phase, which was presumably mediated by Ia inhibitory interneurons. However, it had no effect on H(max)/M(max). Our results indicated that patterned sensory nerve stimulation could modulate the activity of Ia inhibitory interneurons, and this change may have been caused by the synaptic modification of Ia inhibitory interneuron terminals. These results may lead to a clearer understanding of the spinal cord synaptic plasticity produced by repetitive sensory inputs.

Motor training induces plastic changes in the primary motor cortex (M1). However, it is unclear whether and how the latency of motor-evoked potentials (MEP) and MEP amplitude are affected by implicit and/or explicit motor learning. Here, we investigated the changes in M1 excitability and MEP latency induced by implicit and explicit motor learning. The subjects performed a serial reaction time task (SRTT) with their five fingers. In this task, visual cues were lit up sequentially along with a predetermined order. Through training, the subjects learned the order of sequence implicitly and explicitly. Before and after the SRTT, we recorded MEP at 25 stimulation points around the hot spot for the flexor pollicis brevis (FPB) muscle. Although no changes in MEP amplitude were observed in either session, we found increases in MEP latency and changes in histogram of MEP latency after implicit learning. Our results suggest that reorganization across the motor cortices occurs during the acquisition of implicit knowledge. In contrast, acquisition of explicit knowledge does not appear to induce the reorganization based on the measures we recorded. The fact that the above mentioned increases in MEP latency occurred without any alterations in MEP amplitude suggests that learning has different effects on different physiological signals. In conclusion, our results propose that analyzing a combination of some indices of M1 excitability, such as MEP amplitude and MEP latency, is encouraged in order to understand plasticity across motor cortices.

The effects of psychological pressure on corticospinal excitability and muscular activity were investigated in a two-choice reaction task that involved voluntary right index finger movements. In order to induce pressure, participants were given instructions that combined a performance-contingent cash reward, a penalty, and performance comparisons. Following practice, 11 participants performed 40 non-pressure and 40 pressure trials. Results showed successful stress induction, as indexed by significant increases in state anxiety, pulse rate, and galvanic skin response under pressure. Significant increases in the amplitude of the motor evoked potential of the right abductor pollicis brevis (APB) occurred under pressure. The maximum EMG amplitudes of APB and the first dorsal interosseous (FDI) muscle also increased significantly under pressure. These results suggest that descending excitatory volleys in the corticospinal motor tract during the motor preparation processes and peripheral motor unit activities i...

Introduction We investigated the extent to which the corticospinal inputs delivered to Ia inhibitory interneurons influence the strength of disynaptic reciprocal Ia inhibition. Methods Seventeen healthy subjects participated in this study. The degree of reciprocal Ia inhibition was determined via short-latency (condition-test interval: 1–3ms) suppression of Sol H-reflex by conditioning stimulation of common peroneal nerve. The effect of corticospinal descending inputs on Ia inhibitory interneurons was assessed by evaluating the conditioning effect of transcranial magnetic stimulation (TMS) on the Sol H-reflex. Then, we determined the relationship between the degree of reciprocal Ia inhibition and the conditioning effect of TMS on the Sol H-reflex. Result We found that the degree of reciprocal Ia inhibition and the extent of change in the amplitude of the TMS-conditioned H-reflex, which was measured from short latency facilitation to inhibition, displayed a strong correlation ( r =0.76, p Conclusion The extent of reciprocal Ia inhibition is affected by the corticospinal descending inputs delivered to Ia inhibitory interneurons, which might explain the inter-individual variations in reciprocal Ia inhibition.

In primates, unilateral voluntary movements are preferentially controlled by the primary motor cortex (M1) contralateral to the side performing the movement. However, it has been reported that the M1 ipsilateral to the side performing the movement (ipsi-M1) is also activated during unilateral voluntary movements. Recently, studies involving transcranial magnetic stimulation (TMS) or functional magnetic resonance imaging (fMRI) techniques have gradually elucidated the neural mechanisms responsible for modulating ipsi-M1 activity. In particular, the modulation of ipsi-M1 activity is likely to occur in a task-dependent manner, and is also closely associated with advancing age. In addition, ipsi-M1 excitability is suppressed during the acquisition phase of motor learning. Previous studies have suggested that the modulation of ipsi-M1 activity occurs via changes in the activation of the corpus callosum pathways linking the bilateral M1. In this article, we will broadly review the features of ipsi-M1 activity, observed during the execution of unilateral movements, as well as the detailed neural mechanisms underlying the modulation of ipsi-M1 activity. Understanding the role played by ipsi-M1 activity during voluntary movements would improve our knowledge of human motor control systems.

The aim of this study was to investigate the neural mechanisms underlying the changes in the ipsilateral primary motor cortex (ipsi-M1) excitability induced during the unilateral rhythmic muscle contraction of the first dorsal interosseous (FDI) (rhythmic contraction) muscle with three different frequencies of auditory cues (1, 2, and 3 Hz). The effect of different frequencies of unilateral rhythmic contraction on changes in the ipsi-M1 excitability was assessed using a single-pulse transcranial magnetic stimulation (TMS) technique when subjects were performing the unilateral rhythmic contractions according to each auditory cue frequency. After that, the changes in short intracortical inhibition (SICI)/facilitation (ICF), long intracortical inhibition (LICI) within the ipsi-M1, and interhemispheric inhibition (IHI), as well as dorsal premotor cortex to M1 (PMd–M1), and dorsolateral prefrontal cortex to M1 (DLPFC-M1) connectivity from the contralateral hemisphere to the ipsi-M1 were assessed using paired-pulse TMS techniques. The motor evoked potentials (MEP) induced in the right FDI were recorded. In the results, the ipsi-M1 excitability induced in response to single-pulse TMS was significantly decreased in the 2 Hz conditions, compared with the 1 Hz and 3 Hz conditions. Furthermore, PMd–M1 connectivity and LICI were significantly modulated depending on the frequency of the unilateral rhythmic contraction. In contrast, the changes in the SICI, ICF, IHI, and DLPFC–M1 were not directly associated with the rhythm frequency. These results suggest that PMd–M1 connectivity and LICI within the ipsi-M1 are likely to preferentially operate to modulate ipsi-M1 excitability during the performance of unilateral rhythmic contraction with different frequencies.

We examined the effects of observation combined with motor imagery (MI) of a skilled hand-motor task on motor cortex excitability, which was assessed by transcranial magnetic stimulation (TMS). Novices and experts at 3-ball cascade juggling (3BCJ) participated in this study. In one trial, the subjects observed a video clip of 3BCJ while imagining performing it. In addition, the subjects also imagined performing 3BCJ without video clip observation. Motor evoked potentials (MEPs) were recorded from the hand muscles that were activated by the task during each trial. In the novices, the MEP amplitude was significantly increased by video clip observation combined with MI. In contrast, MI without video clip observation significantly increased the MEP amplitude of the experts. These results suggest that action observation of 3BCJ increases the ability of novices to make their MI performing the task. Meanwhile, experts use their own motor program to recall their MI of the task.

Previous studies have shown that spinal neural circuits are modulated by motor skill training. However, the effects of task movement speed on changes in spinal neural circuits have not been clarified. The aim of this research was to investigate whether spinal neural circuits were affected by task movement speed. Thirty-eight healthy subjects participated in this study. In experiment 1, the effects of task movement speed on the spinal neural circuits were examined. Eighteen subjects performed a visuomotor task involving ankle muscle slow (nine subjects) or fast (nine subjects) movement speed. Another nine subjects performed a non-visuomotor task (controls) in fast movement speed. The motor task training lasted for 20 min. The amounts of D1 inhibition and reciprocal Ia inhibition were measured using H-relfex condition-test paradigm and recorded before, and at 5, 15, and 30 min after the training session. In experiment 2, using transcranial magnetic stimulation (TMS), the effects of corticospinal descending inputs on the presynaptic inhibitory pathway were examined before and after performing either a visuomotor (eight subjects) or a control task (eight subjects). All measurements were taken under resting conditions. The amount of D1 inhibition increased after the visuomotor task irrespective of movement speed (P < 0.01). The amount of reciprocal Ia inhibition increased with fast movement speed conditioning (P < 0.01), but was unchanged by slow movement speed conditioning. These changes lasted up to 15 min in D1 inhibition and 5 min in reciprocal Ia inhibition after the training session. The control task did not induce changes in D1 inhibition and reciprocal Ia inhibition. The TMS conditioned inhibitory effects of presynaptic inhibitory pathways decreased following visuomotor tasks (P < 0.01). The size of test H-reflex was almost the same size throughout experiments. The results suggest that supraspinal descending inputs for controlling joint movement are responsible for changes in the spinal neural circuits, and that task movement speed is one of the critical factors for inducing plastic changes in reciprocal Ia inhibition.

Introduction A large number of researches have investigated the mechanisms of implicit and explicit motor sequence learning. The functional changes in motor circuits, which involve the primary motor cortex (M1), are thought to play an important role in memory formation of implicit knowledge. However, it is unclear what functions in the motor circuits contribute to formation of implicit knowledge in the M1. The corticospinal neuromuscular system organizes the modular architectures for generating a variety of movements. Therefore, we hypothesized that performance improvement through implicit learning results from reorganization of the modular architectures. Objectives We used transcranial magnetic stimulation (TMS), data-grove system, and principal component analysis (PCA) to test the hypothesis. Materials and methods Seventeen healthy subjects learned a sequence of serial reaction time task (SRTT) implicitly, and explicitly. Before and after a training session, both TMS-evoked finger joint movements and voluntary finger joint movements during the SRTT were recorded. Results PCA extracted a set of principal components (PCs) from TMS-evoked finger joint movements before (pre-PCs) and after (post-PCs) the training session, respectively. To test an idea that post-PCs contain large amount of information on the learned task, we reconstructed the voluntary movements during SRTT in post-training session by linear combination of a selected subset of the PCs. We found that the quality of the movements reconstructed by the post-PCs was superior to that of the pre-PCs in the implicit condition. By contrast, in the explicit condition, there was no significant difference in the reconstruction quality between the pre-PCs and post-PCs. Conclusion Our results suggest that the implicit knowledge is acquired through learning-specific reorganization of the modular architectures in the corticospinal neuromuscular system. On the other hand, the reorganization of the modular architectures does not relate to acquiring the explicit knowledge.

Objective To investigate whether unilateral voluntary movement affects voluntary drive of motor imagery for the contralateral limb. Methods The subjects were asked to maintain the left index-finger movements with different directions (abduction and adduction, 5% of maximum voluntary contraction; MVC) and with different force levels (10% and 25% MVC). Under these conditions, transcranial magnetic stimulation was applied over the left motor cortex to record motor evoked potential (MEP) from the right first dorsal interosseous muscle with or without motor imagery of the right index-finger abduction. Results Voluntary movement of the left index finger with isodirection, but not mirrored direction, to the imagined movement reduced the MEP enhancement induced by motor imagery. MEP was gradually increased depending on increment of the force level on the left side, while the motor imagery-induced MEP enhancement was consequently reduced. Conclusions Enhancement of the motor cortex excitability driven by motor imagery of the contralateral limb is interfered with by isodirection and forceful movement of the ipsilateral limb, which may be due to an increase in the transcallosal inhibitory effects. Significance Using motor imagery as a therapeutic tool, the voluntary movements on the other side of the body should be taken into account.

[Purpose] The aim of this study was to investigate whether the performance of a combination of observation and motor imagery of chopstick use (complex task) increased corticospinal excitability more than the performance of observation alone. [Subjects and Methods] We recruited 10 healthy subjects with no history of neurological diseases. Corticospinal excitability was assessed with the participants seated in front of a computer screen performing three tasks: (1) control, the subjects were instructed to relax; (2) OBS, the subjects were told to observe an action depicted in the video, and (3) OBS + IMG, the subjects were told to imagine performing an action depicted in a video. During tasks (2) and (3), a video was displayed on the computer screen showing the hand of a male subject using chopsticks to move small items of food from one dish to another (first person perspective). Imagery was performed kinesthetically. [Results] The MEP amplitude in the first dorsal interosseous was significantly increased during OBS+IMG relative to that in the control condition, but not that in the OBS condition. The MEP amplitude in the thenar muscles was significantly different between OBS and OBS+IMG. [Conclusion] These results suggest that the combination of observation and motor imagery of a complex task may be more effective than observation alone for motor rehabilitation purposes., The Society of Physical Therapy Science, 23(5), pp.703-706; 2011

The aim of this study was to determine whether the excitatory ipsilateral primary motor cortex (ipsi-M1) is affected by changes in the frequency of rhythmic voluntary contraction of the left first dorsal interosseous (FDI) induced by repetitive abduction of the left index-finger. Transcranial magnetic stimulations were delivered to the left M1 during repetitive left index-finger abduction at 1, 2, and 3Hz, and motor evoked potentials (MEPs) were simultaneously evoked in the resting right (Rt)-FDI, Rt-abductor pollicis brevis, and Rt-abductor digiti minimi. The stimulus-response (S-R) curve of the MEP at each frequency was recorded. In addition, F-waves were recorded from the Rt-FDI during these rhythmic contraction tasks in order to examine the changes in spinal motoneuron excitability. MEPs were markedly increased under the 3Hz conditions compared with the other conditions. However, F-waves were hardly changed under these conditions. The S-R curve of the MEP induced under the 3Hz conditions was significantly steeper than the curves produced under other conditions. Our results indicate that the excitability of ipsi-M1 is affected by the frequency of rhythmic voluntary contraction of unilateral finger movement, which may be caused by neural inputs delivered via a transcallosal pathway.

Objectives The aim of this study was to investigate the mechanism of modulation of soleus H-reflexes during sustained passive rotation of the hip joint. Methods Healthy men participated as subjects in this study. In the supine position, soleus maximum H-reflexes were recorded during rotation of the hip joint. To examine the roles of muscle spindles, we recorded H-reflexes while applying vibration to muscles around the hip joint. Furthermore, to assess the modulation of pre- and postsynaptic inputs to soleus motoneurons, H-reflexes were conditioned by common peroneal nerve stimulation or transcranial magnetic stimulation (TMS). Results The ratio of the maximum H-reflex to the maximum M-response was significantly reduced in externally rotated positions of the hip joint compared to neutral position. In these positions, presynaptic inhibition, assessed by D1 inhibition, was significantly increased. H-reflexes were reduced during vibration, irrespective of muscles involved and hip position. Facilitation of H-reflex induced by conditioning-TMS showed insignificant differences across all hip positions. Conclusions These findings suggest that Ia afferents induced by rotation of the hip joint presumably inhibit soleus H-reflexes presynaptically. Significance Based on the results, therapists could efficiently modulate the spasticity of the lower legs of hemiplegic patients during rotation of the hip joint.

The present study aimed to further investigate whether the intracortical neural circuits within the primary motor cortex (M1) are modulated during ipsilateral voluntary finger movements. Single- and paired-pulse (interstimulus intervals, ISIs; 3 ms and 12 ms) transcranial magnetic stimulations of the left M1 were applied to elicit motor evoked potential (MEP) in the right first dorsal interosseous (Rt-FDI) muscle during voluntary contractions (10% and 30% maximum voluntary contraction) of the left FDI (Lt-FDI) muscle. F-waves of Rt-FDI muscle were recorded under these left index-finger conditions for ensuring that the excitability changes occur at the supraspinal level. MEPs were also recorded during motor imagery of the left index-finger abduction instead of overt movement. The results showed that, in single-pulse transcranial magnetic stimulation (TMS) paradigm, MEPs in Rt-FDI muscle were markedly enhanced during voluntary contractions of Lt-FDI muscle compared with the complete resting state. In paired-pulse TMS paradigm, the short intracortical inhibition was significantly reduced in proportion to increments of the ipsilateral muscle contraction, whereas the intracortical facilitation had no change. F-wave of Rt-FDI muscle was unchanged under these conditions, while MEP in Rt-FDI muscle was also enhanced during motor imagery of the left index-finger abduction. Based on the present results, it is suggested that the intracortical inhibitory neural circuits may be modulated in the transition from rest to activity of the ipsilateral homonymous muscle. The excitability changes in M1 might be induced by overflows of voluntary drive given to the ipsilateral limb, probably via the transcallosal pathway.

To investigate whether the facilitatory effects of motor imagery (MI) are dependent on motor strategies that vary with posture, we used transcranial magnetic stimulation to examine the effects of two forearm positions on motor-evoked potentials during an MI of index-finger abduction. MI-enhanced motor-evoked potentials of the first dorsal interosseous (prime mover) muscle in the forearm prone position were larger than those in the forearm neutral position. The opposite effects were seen in the extensor carpi radialis (synergist) muscle. These effects correspond to the different electromyography activities in the muscles when performing the actual movements in these two forearm positions. It is suggested that MI reflects different motor strategies in the contribution of agonist and synergist muscles towards a motor task.

To explore the effect of mirror box therapy based on the mirror neuron (MN) system of the primary motor cortex (M1), we examined if direct (without a mirror) and indirect (with a mirror) observation of self-movement in healthy subjects induced changes in motor evoked potential (MEP) evoked by transcranial magnetic stimulation (TMS). MEPs were elicited from the first dorsal interosseous (FDI) and the flexor carpi radialis (FCR) muscles. Somatosensory evoked potentials (SEPs) during self-movement observation were also recorded. Both observations of self-movement with and without a mirror increased MEP amplitude. In addition, increase in MEP amplitude was specific to the prime mover muscle involved in the observed movement. The SEPs increased similar to the MEPs during both observations of self-movement with and without a mirror. We conclude that although the MN system can be activated by observing self-movement in a manner similar to that achieved by observing movement of another person, there were no detectable effect on corticospinal excitability that were specific to movements observed with a mirror.

In the present study, we used transcranial magnetic stimulation (TMS) to investigate the changes in the excitability of the left primary motor cortex (M1) innervating the hand muscles and in short-interval intracortical inhibition (SICI) during speech describing hand or leg movements. In experiment 1, we investigated the effects of the contents of speech on the amplitude of the motor evoked potentials (MEPs) induced during reading aloud and silent reading. In experiment 2, we repeated experiment 1 with an additional condition, the non-vocal oral movement (No-Voc OM) condition, and investigated the change in SICI induced in each condition using the paired TMS paradigm. The MEP observed in the reading aloud and No-Voc OM conditions exhibited significantly greater amplitudes than those seen in the silent reading conditions, irrespective of the content of the sentences spoken by the subjects or the timing of the TMS. There were no significant differences in SICI between the experimental conditions. Our findings suggest that the increased excitability of the left M1 hand area detected during speech was mainly caused by speech-related oral movements and the activation of language processing-related brain functions. The increased left M1 excitability was probably also mediated by neural mechanisms other than reduced SICI; i.e., disinhibition.

We aimed to investigate the effects of the tactile stimulation to an observer's fingertips at the moment that they saw an object being pinched by another person on the excitability of observer's primary motor cortex (M1) using transcranial magnetic stimulation (TMS). In addition, the above effects were also examined during action observation combined with the motor imagery. Motor evoked potentials (MEP) were evoked from the subjects' right first dorsal interosseous (FDI) and abductor digiti minimi (ADM) muscles. Electrical stimulation (ES) inducing tactile sensation was delivered to the subjects' first and second fingertips at the moment of pinching action performed by another person. Although neither the ES nor action observation alone had significant effects on the MEP amplitude of the FDI or ADM, the FDI MEP amplitude which acts as the prime mover during pinching was reduced when ES and action observation were combined; however, no such changes were seen in the ADM. Conversely, that reduced FDI MEP amplitude was increased during the motor imagery. These results indicated that the M1 excitability during the action observation of pinching action combined with motor imagery could be enhanced by the tactile stimulation delivered to the observer's fingertips at the moment corresponding to the pinching being observed.

We examined here the effect of the elbow joint posture on elbow flexor fatigability and strength during isometric contraction. Nine healthy subjects, who gave informed consent, participated in the present study. The subjects were seated in a chair with the shoulder joint at 0°, and the shoulders, pelvis, and thigh were immobilized by straps. The elbow joint posture was positioned at 30, 60, 90 and 120° of elbow flexion. First, subjects produced maximal elbow flexion as a maximum voluntary contraction (MVC) in each elbow joint posture. A myodynamometer was used to measure muscular strength. Secondly, the subjects performed 50% MVC effort for over 60 seconds with visual feedback. The surface electromyograms of the biceps brachii and brachioradialis muscles were recorded. The muscle fatigue was characterized by median power frequency (MDPF) calculated from the power spectrum of surface electromyograms using autoregression models. MDPF were calculated for each section, then 60 seconds was divided into each six sections consisting of 10 seconds. Findings of the present study are: 1) the elbow joint posture requiring the maximum strength is 90°; and 2) MDPF decreased over time in both muscles, and the rate of reduction tended to increase with flexion angle. These findings suggest that the elbow joint posture that demonstrates the maximum muscular strength differs from the one which does not cause muscle fatigue easily. When therapist determines training posture, it is necessary to take this point into consideration.

The aim of this study was to investigate the plasticity of M1 innervating the tibialis anterior muscle (TA) induced by the long-term practice of football juggling using a transcranial magnetic stimulation (TMS) technique. Ten football juggling experts and ten novices participated in this study. Motor evoked potentials (MEP) and the H-reflex were recorded from the right TA during isometric dorsiflexion at 10% of maximum voluntary contraction. The MEP input-output curve of the experts was steeper than that of the novices, and reduced short-interval intracortical inhibition and long-interval intracortical inhibition were observed in the experts. In contrast, the ratio of Hmax to Mmax did not differ between the groups. Our results show that football juggling experts displayed enhanced excitability in the M1 innervating the TA, which was induced by the long-term practice of the ankle movements required to perform football juggling well.

Higashi T, Funase K, Kusano K, Tabira T, Harada N, Sakakibara A, Yoshimura T. Motoneuron pool excitability of hemiplegic patients: assessing recovery stages by using H-reflex and M response. Arch Phys Med Rehabil 2001;82:1604-10. Objectives: To compare the excitability of the motoneuron pools of both the spastic and the unimpaired sides of patients with hemiplegia with a new method by using H-reflexes and M responses. The method determines the ratio of the developmental slope of the H-reflex (Hslp) to the slope of the M response (Mslp). We also examined the relation between the Brunnstrom stages and the Hslp/Mslp. Design: Experiment. Setting: Electrophysiologic experimental laboratory in a Japanese medical school. Patients: Fifteen hemiplegic patients (9 men, 6 women; age range, 48–71yr; mean, 60yr) with spasticity caused by a stroke. Interventions: The subject was instructed to relax while seated in a reclining chair with his foot fixed on an immobile pedal. After the soleus H-reflex and M responses on one side were recorded, the same experimental procedures were carried out on the other side. Main Outcome Measures: Hslp/Mslp Brunnstrom stages. Results: Hslp/Mslp had better predictive value than conventional indicators of motoneuron pool excitability. Hslp/Mslp appeared to be a better match for the bell-shaped pattern of the Brunnstrom stages. Conclusion: Hslp/Mslp is the preferred index for evaluating the motoneuron pool excitability of the spastic side of hemiplegic patients. © 2001 by the American Congress of Rehabilitation Medicine and the American Academy of Physical Medicine and Rehabilitation

In the present study, we examined the electromyogram (EMG) patterns of the soleus and medial gastrocnemius (MG) muscles during rhythmical, two-legged hopping to investigate the contributions of the monosynaptic short- and long-latency stretch reflexes during such a natural movement in human. During rhythmical hopping, soleus muscle is activated reflexly at near-monosynaptic latency by stretch resulting from passive ankle flexion upon landing. Soleus muscle also contracts voluntarily in order to launch the body into the next hop. This is part of the rhythmical bursts of activity producing the hops. Depending on the hopping interval, this phase of activation can follow the short-latency phase or precede landing at very short hopping intervals. In MG, there is an initial phase of activity that stiffens the muscle in preparation for landing, and continues through the contact phase. The monosynaptic reflex response to landing is usually superimposed on this activity. Depending on the hopping interval, both of these responses may be overlaid with activity that is time-locked to the take-off into the next hop, and serves to launch the body into the next hop. However, no evidence for a long-latency stretch reflex was found. In addition, the preferred hopping frequency for all subjects was about 2 Hz. This frequency is associated with a pattern of EMG activity the timing of which indicates that it balances the requirement for a comfortable landing from a hop with the optimal muscle activation required for launching the following hop.

H reflexes were evoked in human soleus by stimulating the tibial nerve at a constant intensity. Each trial was then assigned to one of three groups on the basis of the amplitude of its H reflex; all trials in each group were then full-wave rectified and reaveraged. There was a strong positive relationship between the amplitude of the H reflex and the level of electromyographic activity in the muscle at the time of onset of the H reflex, which reflects the activity of the motoneuronal pool when the afferent volley arrived. Thus, much of the variability of the H reflex is due to small changes in the level of activation of the motoneuronal pool during repeated trials. The steady torque preceding the H reflex was a poor predictor of the H-reflex amplitude, presumably because of the delay between the changes in the electrical activity of motoneurons and the mechanical outcome thereof.

We examined changes in somatosensory evoked potentials (SEPs) during voluntary movement of fingers innervated by the stimulated nerve and those not innervated by the stimulated nerve and the relationship to the kind of movement modality. Analysis showed that the amplitude of most components at F3, C3', and P3, except for P45 at C3, N35 and P45 at P3, decreased during voluntary finger movement tasks. Further, we found that the components of P40 at F3, P45 at C3', and N35 at P3 were increased during the voluntary pulling movement of the second and the third digits compared to those during the voluntary pushing movement of the fourth and the fifth digits, whereas all other components were decreased at F3, C3', and P3. We also found that not all components of SEPs were decreased while some SEPs in middle latency were increased. In conclusion, we confirmed the selectivity in attenuation of the SEPs. Moreover, we noted an interesting finding that the selectivity of attenuation of the SEPs was most frequently observed in the N20, P30 (P25 at F3), N35 (N30 at F3), and P45 (P40 at F3) components at F3, C3', and P3.

The aim of the present study was to investigate whether there is a functional difference in short-latency (SIHI) and long-latency (LIHI) interhemispheric inhibition from the active to the resting primary motor cortex (M1) with paired-pulse transcranial magnetic stimulation during a unilateral muscle contraction. In nine healthy right-handed participants, IHI was tested from the dominant to the nondominant M1 and vice versa under resting conditions or during performance of a sustained unilateral muscle contraction with the right or left first dorsal interosseous muscle at 10% and 30% maximum voluntary contraction. To obtain measurements of SIHI and LIHI, a conditioning stimulus (CS) was applied over the M1 contralateral to the muscle contraction, followed by a test stimulus over the M1 ipsilateral to the muscle contraction at short (10 ms) and long (40 ms) interstimulus intervals. We used four CS intensities to investigate SIHI and LIHI from the active to the resting M1 systematically. The amount of IHI during the unilateral muscle contractions showed a significant difference between SIHI and LIHI, but the amount of IHI during the resting condition did not. In particular, SIHI during the muscle contractions, but not LIHI, significantly increased with increase in CS intensity compared with the resting condition. Laterality of IHI was not detected in any of the experimental conditions. The present study provides novel evidence that a functional difference between SIHI and LIHI from the active to the resting M1 exists during unilateral muscle contractions.

Whether a cortical drive to one limb modulates interhemispheric inhibition (IHI) from the active targeting to the non-active motor cortex (M1) remained unclear. The present study using a conditioning-test transcranial magnetic stimulation (TMS) paradigm aimed to directly demonstrate the modulation of IHI during unilateral voluntary or imagined movement in humans. Subjects were asked to actually perform right index-finger abduction (10–70 % of the maximum voluntary contraction) or to imagine the movement. Conditioning and test TMS with an interstimulus interval of 5, 10, and 15 ms were applied over the left and right M1, respectively, and the test motor evoked potential (MEP) was recorded from the left first dorsal interosseous (FDI) muscle. The conditioning TMS intensity was adjusted ranging from 0.6 to 1.4 (in 0.2 steps) times the resting motor threshold (rMT). With test TMS alone, MEP in the left FDI muscle significantly increased during voluntary or imagined movement of the right index-finger. MEP amplitude was significantly reduced in proportion to increments of the conditioning TMS intensity at rest (1.2 and 1.4 times the rMT, P

The effect of performance of a sensorimotor task on the interhemispheric inhibition (IHI) induced from the active primary motor cortex (M1) to the resting M1 was examined in 10 right-handed subjects. Transcranial magnetic stimulation (TMS) was performed to produce motor evoked potentials (MEP) in the resting right (Rt)-first dorsal interosseous (FDI). For the paired-TMS paradigm, a conditioning stimulus (CS) was delivered to the Rt-M1, and its intensity was adjusted from 0.6 to 1.4 times the resting motor threshold of the MEP in the left (Lt)-FDI in 0.2 steps. The test stimulus was delivered to the Lt-M1, and its intensity was adjusted to evoke similar MEP amplitudes in the Rt-FDI among the task conditions. The interstimulus interval was fixed at 10 ms. As a sensorimotor task, a fine-motor manipulation (FM) task (using chopsticks to pick up, transport, and release glass balls) was adopted. In addition, an isometric abduction (IA) task was also performed as a control task. These tasks were carried out with the left hand. The IHI from the active to the resting M1 observed during the FM task was markedly increased compared with that induced during the IA task, and this effect was not dependent on the MEP amplitude evoked in the active Lt-FDI by the CS. The present findings suggest that the increased IHI from the active to the resting M1 observed during the FM task was linked to reductions in the activity of the ipsilateral intracortical inhibitory circuit, as we reported previously.

The 1st purpose of this study was to examine multiple electromyography (EMG) during voluntary hand movements. A secondary purpose was to investigate possible effects of pressure on intracortical inhibition (ICI) and intracortical facilitation (ICF) functions of the motor cortex, using paired-pulse transcranial magnetic stimulation. Twelve participants traced a 15-cm diameter target circle using a small laser pointer attached to the right index finger. After 5 acquisition trials, they performed 3 nonpressure trials followed by 3 pressure trials. The results showed that pressure had effects not only on agonist EMG activity but also on multiple muscles, such as synergist. In addition, a decrease in ICI and an increase in ICF were both observed under pressure for muscles other than the agonist.

We reviewed the literature on basic psychological correlates of the well-known phenomenon of the location-distance interference in motor short-term memory (Kerr, 1978 ; Walsh, Russell, Imanaka, & James, 1979). The location-distance interference in motor short-term memory has frequently been demonstrated as an unavoidable interference phenomenon observed in the reproduction of movement location and distance in arm positioning. The most important aspect of this phenomenon is that even when a subject concentrates on a specific cue (i.e., either end-location or distance) the other cue is also coded unintentionally and, as a result, the reproduction movement guided on the basis of the specific cue is unavoidably influenced by the other nonspecific cue. In thls review article, we first reviewed the literature on the basic theories and nature of shortterm memory, particularly on the limited processing capacity. We then referred to the unlimited, automatic processing in visual-verbal domains, referring to the Stroop phenomenon. Finally, in conjunction with the notion of automatic processing, we examined the possible aspects of information processing which may be responsible for mediating the location-distance interference in motor short-term memory, The Annals of physiological anthropology, 12(5), pp.269-283; 1993

The intracellular biochemical process underlying oxytocin-induced change of membrane properties was analyzed in an identified neuron of Achatina fulica Férussac, using pressure injection technique and pharmacological tools. Oxytocin dose-dependently enhanced the negative slope resistance (NSR) region on the current-voltage relation. The oxytocin-induced current was attenuated by a reduction of extracellular Na+ and not influenced by the addition of 100 microM tetrodotoxin (TTX) to the medium, suggesting that this current is predominantly due to the activation of TTX-resistant Na+ channels. In the Ca2(+)-free state, substituted by an equivalent amount of Co2+, the amplitude of oxytocin-induced current was somewhat reduced at the NSR region but it was not influenced at less than -60 mV. Application of 100 microM isobutylmethylxanthine, a phosphodiesterase inhibitor, augmented the oxytocin-induced current. Pressure injection of 10 mM adenosine 3',5'-cyclic monophosphate (cAMP) elicited a Na(+)-dependent inward current similar to the oxytocin response. The further role of cAMP linked with the oxytocin-induced current was investigated using two kinds of cAMP-dependent protein kinase inhibitors, isoquinolinesulfonamide (H-8) and protein kinase inhibitor (PKI). Extracellular application of H-8 or pressure injection of PKI, prior to oxytocin application, both blocked the oxytocin-induced current. Based on these results, oxytocin-elicited inward currents may mediate cAMP-dependent protein phosphorylation mainly by activation of Na+ channels.

Although there were many reports relating to intermanual transfer of behavioral motor tasks in humans, it is still not well-known whether the transfer phenomenon between the trained and untrained hand is accompanied by corresponding changes in motor system. In the present study we applied transcranial magnetic stimulation to investigate the practice effects of unilateral fingertip precision grip on corticospinal excitability, regarding both the trained and untrained hand muscles. The results showed that after practice fingertip grip force became steady and safety margin dramatically decreased not only in the trained hand, but also in the untrained hand. Regarding MEP and background EMG (B.EMG) activities, the regression slope of MEP/B.EMG ratio in the first dorsal interosseous (FDI) muscle became significantly steeper after practice in both hands, but in the thenar (TH) muscle there were no clear modulations. These results indicated that through practice qualitative or functional changes of corticospinal systems related to the reorganization for a fingertip precision grip prominently reflect only on FDI muscle which plays a dominant role in the task. More importantly, such effects were simultaneously seen in the untrained hand correspondent to the trained hand, i.e., changes of input-output property in M1 occur not only in the trained hand, but also in the untrained hand. Based on the present results, we suggest that training-induced neural adaptations of the central nervous system may include improvement of its predicting fingertip grip force for self-lifting of the object in the untrained hand.