Temporal modulations of agonist and antagonist muscle activities accompanying improved performance of ballistic movements.

Although many studies have examined performance improvements of ballistic movement through practice, it is still unclear how performance advances while maintaining maximum velocity, and how the accompanying triphasic electromyographic (EMG) activity is modified. The present study focused on the changes in triphasic EMG activity, i.e., the first agonist burst (AG1), the second agonist burst (AG2), and the antagonist burst (ANT), that accompanied decreases in movement time and error. Twelve healthy volunteers performed 100 ballistic wrist flexion movements in ten 10-trial sessions under the instruction to "maintain maximum velocity throughout the experiment and to stop the limb at the target as fast and accurately as possible". Kinematic parameters (position and velocity) and triphasic EMG activities from the agonist (flexor carpi radialis) and antagonist (extensor carpi radialis) muscles were recorded. Comparison of the results obtained from the first and the last 10 trials, revealed that movement time, movement error, and variability of amplitudes reduced with practice, and that maximum velocity and time to maximum velocity remained constant. EMG activities showed that AG1 and AG2 durations were reduced, whereas ANT duration did not change. Additionally, ANT and AG2 latencies were reduced. Integrated EMG of AG1 was significantly reduced as well. Analysis of the alpha angle (an index of the rate of recruitment of the motoneurons) showed that there was no change in either AG1 or AG2. Correlation analysis of alpha angles between these two bursts further revealed that the close relationship of AG1 and AG2 was kept constant through practice. These findings led to the conclusion that performance improvement in ballistic movement is mainly due to the temporal modulations of agonist and antagonist muscle activities when maximum velocity is kept constant. Presumably, a specific strategy is consistently applied during practice.

Preparatory suppression of the human primary motor cortex induced by repetition of simple and choice reaction time tasks: a transcranial magnetic stimulation study.

Transcranial magnetic stimulation was performed to investigate preparatory suppression of activity in the human primary motor cortex (M1) in relation to trial repetition of simple (SRT) and Go/NoGo choice RT (CRT) tasks. These tasks were performed in such a way that after a warning signal, the subjects (N=16) maintained 5% MVC isometric finger force against the force sensor to secure a facilitated state of M1. A response signal to generate pulsed force came at 2 s after the warning signal. TMS was given 1.5 s after the warning signal, and the amplitudes of motor-evoked potentials (MEPs) in the first dorsal interosseous muscle were evaluated during 30 repetitive trials over 3 sessions for each subject. For the SRT task, the MEP amplitude was significantly decreased from baseline values in all trials of the three sessions. For the CRT task, on the other hand, there was a clear decreasing trend of the MEP amplitude with trial at the first and second sessions. The mean MEP amplitude at the first session was clearly higher than the baseline while it decreased significantly and reached the value below the baseline at the third session. The findings indicate that active suppression of M1 activity is involved in the preparatory state for RT tasks and that the degree of this suppression can relate to trial experience. The effect is thus most likely a consequence of a rapid adaptive change with the central nervous system in optimizing the preparatory state of M1 for the upcoming motor response.

Effects of motor imagery are dependent on motor strategies.

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.

Effects of intermanual transfer induced by repetitive precision grip on input-output properties of untrained contralateral limb muscles.

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.

Motor strategies and excitability changes of human hand motor area are dependent on different voluntary drives.

The present study examined whether there were different voluntary drives between intended and non-intended muscle contractions. In experiment 1, during intended and non-intended muscle contractions, electromyograms (EMGs) were recorded from the first dorsal interosseous (FDI) and extensor carpi radialis (ECR) muscles when force levels were varied from 10% to 50% maximal voluntary contraction (MVC) in 10% MVC steps. In experiment 2, using transcranial magnetic stimulation, motor-evoked potentials (MEPs) were recorded from the FDI muscle when EMGs were varied from 10% to 40% EMGmax (EMG activities during MVC) in 10% EMGmax steps during intended and non-intended muscle contractions. In experiment 3, at 10% MVC force level MEPs were recorded before and after practice. The results showed that, in the FDI muscle, EMGs during intended muscle contractions were larger than those during non-intended ones at higher force levels (30-50% MVC). In the ECR muscle, reverse results were observed. At comparable EMG levels of the FDI muscle MEPs were the same during intended and non-intended muscle contractions. After practice, MEPs during intended muscle contraction became larger than those during non-intended at 10% MVC force level, while EMGs were the same between two muscle contractions. It is concluded that motor strategies and excitability changes of hand motor area are different during intended and non-intended muscle contractions, and these differences are due to the different voluntary drives of intended and non-intended. The present findings may contribute to the understanding of rehabilitation for patients suffering from damages of the central motor system.

Excitability changes in human hand motor area induced by voluntary teeth clenching are dependent on muscle properties.

To investigate whether the early effects of voluntary teeth clenching (VTC) among the first dorsal interosseous (FDI), abductor digiti minimi (ADM), and abductor pollicis brevis (APB) muscles are differently modulated depending on their muscle properties, we examined the responses of motor evoked potentials (MEPs) elicited by transcranial magnetic stimulation with selected current directions and by brainstem magnetic stimulation (BMS). Although MEP responses with anterior-medially current direction (preferentially elicited I1-waves) were facilitated in all three muscles, those responses with posterior-laterally current direction (preferentially elicited I3-waves) were different among FDI, ADM, and APB muscles. That is, MEP responses in FDI and APB muscles were significantly reduced, whereas those responses in ADM muscle were not significantly reduced. Further, inhibitory effects of VTC in FDI muscle were more potent than those in ADM or APB muscles. On the other hand, the responses to BMS were unchanged by VTC in all three muscles, suggesting that the modulations of MEP were attributed to the cortical origin. On the basis of our previous findings that the inhibitory connections in FDI muscle are more potent than those in ADM muscle (Takahashi et al. in Clin Neurophysiol 116:2757-2764, 2005), the cortical effects of VTC among three hand muscles are differently modulated, depending on muscle properties, presumably the extents of inhibitory connections to corticospinal tract neurons. Considering that the functional capacity in FDI muscle is higher than that in ADM or APB muscles, the cortical inhibitory effect of VTC might contribute to the sophisticated regulation of the motor outputs even during VTC.

Functional demanded excitability changes of human hand motor area.

The present study was performed to examine if there are functional differences between the first dorsal interosseous (FDI) and the abductor digit minimi (ADM) muscles during different muscle contractions, namely dynamic and static contractions of the index and little finger abductions. It was also examined whether these functional differences occur at the cortical level. The motor evoked potentials (MEPs) elicited by transcranial magnetic stimulation (TMS) and force curves, during the muscle contractions, were simultaneously recorded. Rest motor thresholds (RMTs) and active motor thresholds (AMTs), during dynamic and static contractions, were determined in the two muscles. In all trials, the background EMGs (B.EMGs) were kept at the same level in each muscle. Results showed that the target matching errors of dynamic contractions were statistically smaller in the FDI muscle than those in the ADM. In the FDI muscle, the AMT during dynamic contractions was significantly lower than during static ones and the MEPs elicited by TMS were larger during dynamic contractions than those during static ones. However, such results were not found in the ADM muscle. In order to investigate whether the differences were caused by the excitability changes that occurred in the cortical level, the responses elicited by subcortical stimulations were recorded using the same procedures as the experiment of TMS. Responses to subcortical stimulations during dynamic contractions were similar to those during static ones in either muscle. It is concluded that there are differences in the task-dependent MEP facilitations between the FDI and ADM muscles. And the differences are due to the functional demanded excitability changes accompanied by the cortical activation.

Differential modulations of intracortical neural circuits between two intrinsic hand muscles.

OBJECTIVE: To investigate whether the intracortical inhibitory (ICI) and facilitatory (ICF) circuits in the primary motor cortex between the first dorsal interosseous (FDI) and abductor digiti minimi (ADM) muscles are modulated differently. METHODS: We conducted paired-pulse transcranial magnetic stimulation in combination with different current directions (anterior-medially: AM, and posterior-laterally: PL) under relaxed and active muscle conditions with interstimulus intervals (ISIs) between 2 and 16 ms. RESULTS: In both muscle conditions, the conditioned motor-evoked potential (MEP) responses obtained with the AM current direction (preferentially eliciting early I-waves) were similar between the two muscles at all ISIs, but the MEP responses obtained with the PL current direction (preferentially eliciting late I-waves) were different between FDI and ADM muscles, in that the conditioned MEP responses in FDI muscle were inhibited at all ISIs under both muscle conditions, whereas those in ADM muscle were suppressed at only short ISIs (2-4 ms). CONCLUSIONS: These results indicate that the inhibitory connections operating for the corticospinal tract neurons in FDI muscle are more potent, and, conversely, that those in ADM muscle are weaker. SIGNIFICANCE: The different modulations of ICI circuits between FDI and ADM muscles is an important neural mechanism which may contribute to different functional demands (finger dexterity).

Physical practice induces excitability changes in human hand motor area during motor imagery.

The present study was undertaken to investigate the effects of physical practice on excitability changes in human primary motor cortex (M1) during motor imagery (MI). Using different intensities of transcranial magnetic stimulation (TMS), we examined changes in the motor evoked potential (MEP) of the first dorsal interosseous (FDI) muscle with and without MI, and before and after physical practice. On comparing results for MEPs recorded before and after physical practice, the difference between the MEP amplitudes observed at rest and during MI only increased at higher TMS intensities. This finding indicates a physical practice-dependent increase of the higher threshold recruitment of corticospinal tract neurons (CTNs), consistent with synchronization for efficient movement, and provides evidence that neural mechanisms of MI depend not only on the type of movement but also on the extent of the motor adaptation (the physical practice). These present findings also show the benefit of MI and highlight beneficial neural mechanisms related to the activation of M1 during MI. In other words, MI may reflect functional changes of M1 that are similar to the changes observed after physical practice.

Modulations of input-output properties of corticospinal tract neurons by repetitive dynamic index finger abductions.

The goal of this study was to investigate how corticospinal tract neurons (CTNs) are modulated after repetitive dynamic muscle contractions. To address this question, changes of motor evoked potentials (MEPs) to transcranial magnetic stimulation and background EMG (B.EMG) activities were examined. Subjects were instructed to perform an isometric dynamic index finger abduction as accurately as possible under the target-force-matching tasks (10% or 30% MVC), while MEPs of a first dorsal interosseous (FDI) were elicited during performance of the task. After repetitive dynamic FDI contractions (100 trials), the following remarkable phenomena were observed: (1) both B.EMG activities and MEP amplitudes decreased in proportion to the number of trials, (2) these phenomena were most commonly observed in different conditions, i.e., different force levels and hands (preferred or non-preferred hands), and (3) after repetition of the tasks, the MEP amplitude/B.EMG (MEP/B.EMG) ratio became smaller. Decreases of B.EMG activities with reduction of MEP amplitudes and diminishing MEP/B.EMG ratio might suggest the occurrence of reorganization of input-output properties in CTNs for an efficient performance as a function of motor adaptation. Thus, we conclude that motor adaptation after repetitive dynamic muscle contractions probably occurs less specifically and due to susceptible modulations of spinal motoneurons reflected in the integrative functions of CTNs.

Excitability changes of motor evoked potentials dependent on muscle properties and contraction modes.

Using transcranial magnetic stimulation (TMS), differences in the excitability changes of motor evoked potentials (MEPs) between isometric (force task) and isotonic (movement task) muscle contractions in a distal (first dorsal interosseous; FDI) and a proximal (middle deltoid; MD) muscle were studied. In the FDI muscle, the active threshold of MEP recruitment was significantly lower in the isotonic than that in the isometric muscle contraction in spite of identical background EMG activity levels. Additionally, the dependence of the MEP amplitude on background EMG activity was significantly greater in the isotonic than in the isometric muscle contraction at low EMG activity levels, but the difference disappeared beyond middle EMG activity levels. In the MD muscle, the dependence of the MEP amplitude on background EMG activity was significantly greater in the isotonic than in the isometric muscle contraction, and further this dependence was kept at all muscle contraction levels. These results indicate that the dependence of the MEP amplitude on background EMG activity is modulated not only by the different muscle contraction modes (isotonic and isometric), but also by muscle properties (distal and proximal). Thus, the present findings suggest that the task-specific extra excitation in the proximal muscle is definitely produced corresponding to task differences (task-dependent subliminal fringe), which might be explained by the predominant frequency principle if applied to the proximal muscle. On the other hand, the lack of task-dependent extra excitation in the distal muscle is explained by the predominant recruitment principle for force grading in small hand muscles.

Effect of vibration-induced postural illusion on anticipatory postural adjustment of voluntary arm movement in standing humans.

We investigated the contribution of sensory signals arising from muscle proprioceptive receptors to anticipatory postural adjustments (APAs). During vibration applied to ankle (tibialis anterior; TA, soleus; Sol) or neck muscles, subjects generally describe having illusory sensations of whole-body movement, namely, whole-body movement in a backward and forward direction induced by vibration of the Sol or TA, respectively, and the front or back surface of the neck muscles, respectively. Preceding electromyographic (EMG) activity of the ipsilateral biceps femoris (BFi) muscle induced by rapid voluntary arm movement and the typical phenomenon of APA were changed dependent on these illusory whole-body movements, with preceding EMG activities of BFi appearing earlier in vibration applied to TA and later in vibration applied to Sol muscle. In vibration applied to the back surface of neck muscle, preceding EMG activities of BFi appeared earlier, as with vibration applied to TA. On the contrary, in vibration applied to the front surface of neck muscles, preceding EMG activities of BFi appeared later, as with vibration applied to Sol. Based on these results, we discuss changes in the central processing of proprioceptive signals used for coding of the spatial orientation of the body and its contribution to postural stabilization.