Constraints on hand-foot coordination associated with phase dependent modulation of corticospinal excitability during motor imagery.

Interlimb coordination involving cyclical movements of hand and foot in the sagittal plane is more difficult when the limbs move in opposite directions compared with the same direction (directional constraint). Here we first investigated whether the directional constraint on hand-foot coordination exists in motor imagery (imagined motion). Participants performed 10 cyclic coordinated movements of right wrist flexion-extension and right ankle dorsiflexion-plantarflexion as quickly and precisely as possible, in the following three conditions; (1) actual movements of the two limbs, (2) imaginary movements of the two limbs, and (3) actual movement of one limb combined with imaginary movement of the other limb. Each condition was performed under two directions; the same and the opposite direction. Task execution duration was measured as the time between the first and second press of a button by the participants. The opposite directional movement took a significantly longer time than did the same directional movement, irrespective of the condition type. This suggests that directional constraint of hand-foot coordination occurs even in motor imagery without actual motor commands or kinesthetic signals. We secondarily examined whether the corticospinal excitability of wrist muscles is modulated in synchronization with an imaginary foot movement to estimate the neural basis of directional constraint on imaginary hand-foot coordination. The corticospinal excitability of the forearm extensor in resting position increased during dorsiflexion and decreased during plantarflexion similarly in both actual and imaginary foot movements. This corticospinal modulation depending on imaginary movement phase likely produces the directional constraint on the imaginary hand-foot coordination.

Stability of bimanual finger tapping coordination is constrained by salient phases.

In bimanual cyclical continuous movements, the relative timing of the most salient movement phase in each movement is a predominant constraint. This is the case for coordination when both movements have a single most salient phase (the relative-salience hypothesis). We tested whether the relative-salience hypothesis could explain results obtained for repetitive discrete movements, utilizing finger tapping. In experiment 1, participants performed unimanual alternate two-finger tapping with the metronome beat (i.e., one finger taps on the beat and the other finger taps off the beat). The stability of the tapping timing relative to the beat, which reflects the extent of salience, was higher in the index finger than the middle finger, and was lower in the ring finger than the middle finger. In experiment 2, participants performed four conditions of repetitive bimanual four-finger tapping (i.e., alternate two-finger tapping in each hand) without external pacing signals. Under all four conditions, a more stable pattern occurred when the timing of the more salient tapping in each hand was simultaneous rather than alternate, regardless of relative direction in the external space or movement coupling of the homologous fingers. The results indicated that bimanual four-finger tapping could be explained by the relative-salience hypothesis.

Reaching movements are automatically redirected to nearby options during target split.

Motor behavior often occurs in environments with multiple goal options that can vary during the ongoing action. We explored this situation by requiring subjects to select between different target options during an ongoing reach. During split trials the original target was replaced with a left and a right flanking target, and participants had to select between them. This contrasted with the standard jump trials, where the original target would be replaced with a single flanking target, left or right. When participants were instructed to follow their natural tendency, they all tended to select the split target nearest the original. The near-target preference was more prominent with increased spatial disparity between the options and when participants could preview the potential options. Moreover, explicit instruction to obtain the "far" target during split trials resulted many errors compared with a "near" instruction, ~50% vs. ~15%. Online reaction times to target change were delayed in split trials compared with jump trials, ~200 ms vs. ~150 ms, but also highly automatic. Trials in which the instructed far target was correctly obtained were delayed by a further ~50 ms, unlike those in which the near target was incorrectly obtained. We also observed nonspecific responses from arm muscles at the jump trial latency during split trials. Taken together, our results indicate that online selection of reach targets is automatically linked to the spatial distribution of the options, though at greater delays than redirecting to a single target.NEW & NOTEWORTHY This work demonstrates that target selection during an ongoing reach is automatically linked to the option nearest a voided target. Online reaction times for two options are longer than redirection to a single option. Attempts to override the near-target tendency result in a high number of errors at the normal delay and further delays when the attempt is successful.

Spinal Circuits Mediate a Stretch Reflex Between the Upper Limbs in Humans.

Inter-limb reflexes play an important role in coordinating behaviors involving different limbs. Previous studies have demonstrated that human elbow muscles express an inter-limb stretch reflex at long-latency (50-100 ms), a timing consistent with a trans-cortical linkage. Here we probe for inter-limb stretch reflexes in the shoulder muscles of human participants. Unexpected torque pulses displaced one or both shoulders while participants adopted a steady posture against background torques. The results demonstrated inter-limb stretch reflexes occurring at short-latency for both shoulder extensors and flexors; the rapid timing (36-50 ms) must involve a spinal linkage for the two arms. Inter-limb stretch reflexes were also observed at long-latency yet they were opposite to the preceding short-latency; when the short-latency stretch reflex was excitatory then the long-latency stretch reflex was inhibitory and vice versa. Comparing the responses to contralateral arm displacement to those during simultaneous displacement of both arms revealed that inhibitory inter-limb stretch reflexes are independent of within-limb stretch reflexes, but that excitatory inter-limb stretch reflexes are suppressed by within-limb stretch reflexes. Our results provide the first demonstration of short-latency inter-limb stretch reflexes in the upper limb of humans and reveal interacting spinal circuits for within-limb and inter-limb stretch reflexes.

Performance evaluation of lysimeter experiments for simulating pesticide dissipation in paddy fields. Part 1: Submerged application of granular pesticides.

Three-year comprehensive experiments were conducted to compare the dissipation patterns of a total of 16 pesticides, including 3 metabolites, as granular formulations applied in lysimeters and paddy fields with two soil types. Analytical concentrations of the target pesticides in paddy water were analyzed using a granular kinetic model consisting of the following parameters: release rate, decrease rate, and dissolved concentration. Results of parameter grouping analyses of the kinetic models showed that 56% of data reproducibility (entire grouping) was obtained between replicates for the lysimeters. In comparisons between the lysimeters and paddy fields, 48% of decrease rates and 34% of release rates were grouped, although significant differences were observed with a nearly 90% difference for dissolved concentrations. These differences might be attributed to the hydrological components such as water management and meteorological covariates in paddy fields, the daily percolation in lysimeters and the adsorption-desorption kinetics between paddy water and soil.

Performance evaluation of lysimeter experiments for simulating pesticide dissipation in paddy fields. Part 2: Nursery-box application and foliar application.

Comparative experiments investigating the dissipation of four nursery-box-applied pesticides and three foliar-applied pesticides were conducted using lysimeters and in actual paddy fields. In the lysimeter experiments, there were test plots for submerged application for both application types. Analytical concentrations of the pesticides in paddy water were evaluated using appropriate kinetic models. The detection levels of pesticides in the paddy water for the nursery-box and foliar applications were 10-77% and 42-79% of the submerged application, respectively. The times required for 50% dissipation (DT 50s) in case of the nursery-box and foliar applications were 0.8-10.4 days and 0.5-2.7 days, respectively. Although overall dissipations were affected by the physicochemical properties of the pesticide and the experimental design in the test plots, the initial detection levels in the lysimeters, governed by the runoff at transplanting and the deposition at spraying, were comparable with those in the actual paddy fields.

Effect of salient points in movements on the constraints in bimanual coordination.

The relative-salience hypothesis has been proposed as a possible explanation for the stability of bimanual coordination. This explanation proceeds from a psychological viewpoint and is based on the following tenets: (1) cyclic joint motions involving two movements are conceived of as a unified event, (2) if a "single" point in each movement is seen as the most salient, the salient points of the two movements prefer to go together, and (3) in other cases, a unified event will be constrained by movement direction. In this investigation, we examined whether the relative-salience hypothesis could predict the type of constraint (i.e., action coupling vs movement direction) for various bimanual coordination movements. Participants performed six different joint movements in synchrony with metronome beats. Both index finger flexion/extension and forearm pronation/supination had a "single" salient point (JMsingleSP), the others had "two" salient points (JMtwoSP). Then, we applied the relative-salience hypothesis to four bimanual coordinations. The coupling of simultaneous forearm pronation was more stable than alternate pronation. Similarly, the coupling of finger flexion and forearm pronation was more stable than that of finger flexion and forearm supination. For the coordination of radial flexion/ulnar flexion and index finger flexion/extension as well as forearm pronation/supination and radial flexion/ulnar flexion, symmetric movements were more stable than asymmetric movements. The results indicated that the stability of bimanual coordination was predominantly constrained by coupling of salient points when using two JMsingleSP and it was predominantly constrained by movement direction when coordinating JMsingleSP and JMtwoSP. Thus, the relative-salience hypothesis was supported.

Muscle Relaxation of the Foot Reduces Corticospinal Excitability of Hand Muscles and Enhances Intracortical Inhibition.

The object of this study was to clarify the effects of foot muscle relaxation on activity in the primary motor cortex (M1) of the hand area. Subjects were asked to volitionally relax the right foot from sustained contraction of either the dorsiflexor (tibialis anterior; TA relaxation) or plantarflexor (soleus; SOL relaxation) in response to an auditory stimulus. Single- and paired-pulse transcranial magnetic stimulation (TMS) was delivered to the hand area of the left M1 at different time intervals before and after the onset of TA or SOL relaxation. Motor evoked potentials (MEPs) were recorded from the right extensor carpi radialis (ECR) and flexor carpi radialis (FCR). MEP amplitudes of ECR and FCR caused by single-pulse TMS temporarily decreased after TA and SOL relaxation onset, respectively, as compared with those of the resting control. Furthermore, short-interval intracortical inhibition (SICI) of ECR evaluated with paired-pulse TMS temporarily increased after TA relaxation onset. Our findings indicate that muscle relaxation of the dorsiflexor reduced corticospinal excitability of the ipsilateral hand muscles. This is most likely caused by an increase in intracortical inhibition.

Corticospinal excitability modulation in resting digit muscles during cyclical movement of the digits of the ipsilateral limb.

We investigated how corticospinal excitability of the resting digit muscles was modulated by the digit movement in the ipsilateral limb. Subjects performed cyclical extension-flexion movements of either the right toes or fingers. To determine whether corticospinal excitability of the resting digit muscles was modulated on the basis of movement direction or action coupling between ipsilateral digits, the right forearm was maintained in either the pronated or supinated position. During the movement, the motor evoked potential (MEP) elicited by transcranial magnetic stimulation (TMS) was measured from either the resting right finger extensor and flexor, or toe extensor and flexor. For both finger and toe muscles, independent of forearm position, MEP amplitude of the flexor was greater during ipsilateral digit flexion as compared to extension, and MEP amplitude of the extensor was greater during ipsilateral digit extension as compared to flexion. An exception was that MEP amplitude of the toe flexor with the supinated forearm did not differ between during finger extension and flexion. These findings suggest that digit movement modulates corticospinal excitability of the digits of the ipsilateral limb such that the same action is preferred. Our results provide evidence for a better understanding of neural interactions between ipsilateral limbs, and may thus contribute to neurorehabilitation after a stroke or incomplete spinal cord injury.

Potential explanation of limb combination performance differences for two-limb coordination tasks.

Rhythmic two-limb coordinated movements in the sagittal plane are variable and inaccurate when the movements are in the opposite direction as compared with those in the same direction (directional constraint). The magnitude of directional constraint depends on the particular limb combination. It is prominent in ipsilateral hand-foot coordination, but minimal in bimanual hand coordination. The reason for such differences remains unclear. In this study, we investigated the possible mechanisms underlying the production of the difference that depend on limb combination. Subjects performed two-limb rhythmic coordinated movements either in the same or in the opposite direction for three separate limb combinations (bilateral hands, contralateral hand and foot, and ipsilateral hand and foot). For each combination two different tasks were performed. In the first condition, subjects actively moved two limbs (active condition). Second, subjects actively moved one limb in coordination with a passively moved limb (passive condition). In the active condition, the directional constraint was dependent upon the limb combination, as reported in previous studies; the directional constraint was quite prominent in ipsilateral combinations, intermediate in contralateral combinations, and minimal for bilateral combination. However, differences in the directional constraint did not depend on limb combination for any combination in the passive conditions which apparently utilized closed-loop control. In other word, the difference depending on limb combination disappeared when control strategies become uniformly closed-loop. Thus, we speculate that the control strategy utilized depends on limb combination in the active condition. Additionally, different mechanisms other than closed-loop control also would have influence depending on the particular limb combination. This may result in differences in performance depending upon the limb combination.

Motor imagery of voluntary muscle relaxation induces temporal reduction of corticospinal excitability.

Voluntary muscle relaxation is an "active process" requiring cortical activation. However, cortical activation during motor imagery of muscle relaxation has not been well understood. The purpose of this study was to clarify time-dependent changes in corticospinal excitability during the imagery of muscle relaxation. Ten participants imagined volitional muscle relaxation from an imagined pinching with their right index finger and thumb in response to an auditory cue. Transcranial magnetic stimulation was applied at the left primary motor area of the first dorsal interosseous (FDI) muscle at different time intervals after the auditory cue. Motor evoked potentials (MEPs) were recorded from the right hand and forearm muscles. The MEP amplitudes of the FDI and the synergist temporally decreased after the auditory cue as compared with those present in the resting condition. Our finding indicates that motor imagery of muscle relaxation induces a temporal reduction of the corticospinal excitability related to the targeted muscle.

Interaction between simultaneous contraction and relaxation in different limbs.

We investigated the interaction between relaxation and contraction for remote, ipsilateral muscles of the hand and foot (wrist extensor/ankle dorsiflexor). Subjects sat in an armchair and were able to freely move their right hand and foot. They performed eight tasks: (1) wrist extension from a flexed (relaxed) position, (2) ankle dorsiflexion from a plantarflexed (relaxed) position, (3) wrist relaxation from an extended position, (4) ankle relaxation from a moderately dorsiflexed position. The remaining tasks involved (5) simultaneous performance of tasks 1 and 2, (6) 3 and 4, (7) 1 and 4, and (8) 2 and 3. Subjects performed each task as fast as possible after hearing a start signal. They were instructed not to activate antagonistic muscles in the tasks involving relaxation. When contraction of the wrist flexor/ankle dorsiflexor was concurrent with relaxation of the ipsilateral limb (ankle dorsiflexor/wrist extensor), the reaction time of contraction as observed in electromyography (EMG) activities in the wrist extensor/ankle dorsiflexor became longer and the EMG activities became smaller. Our findings suggest that muscle relaxation in one limb interferes with smooth movement of muscle contraction in the ipsilateral limb.

Factors that determine directional constraint in ipsilateral hand-foot coordinated movements.

In performing simultaneous rhythmic movements of the ipsilateral hand and foot, there are differences in the level of stability between same directional (stable) and opposite directional (unstable) movements. This is the directional constraint. In this study, we investigated three factors ("interaction in efferent process," "interaction of afferent signals," and "error correction") proposed to underlie for the directional constraint. We compared the performance of three tasks: (1) coordination of actively moved ipsilateral hand and foot, (2) active hand movement in coordination with passively moved foot, (3) active hand movement not coordinated with a passively moved foot. In each task, both same and opposite directional movements were executed. There was no difference between performance estimated with success rate for the first and second task. The directional constraint appeared in both tasks. Thus, the interaction in efferent processes, which was shown to be responsible for the directional constraint in bimanual coordination, was not involved with the directional constraint of ipsilateral hand-foot coordination. The directional constraint did not appear in the third task, which suggested that "interaction of afferent signals" also had no contribution. These results indicated that "error correction" must be the most critical of these factors for mediating the directional constraint in ipsilateral hand-foot coordinated movements.

Brain activity during motor imagery of an action with an object: a functional magnetic resonance imaging study.

We utilized functional magnetic resonance imaging to investigate the brain regions activated during motor imagery of an action with an object both with and without passively holding the object. Participants performed the following tasks: (1) 'Imagery with Ball' condition: subjects imagined squeezing a foam ball (7cm diameter) while holding the ball, (2) 'Imagery' condition: subjects imagined squeezing a ball without holding the ball, and (3) 'Ball' condition: subjects held the ball without motor imagery. Regions activated by the 'Imagery with Ball' condition were located in the left dorsolateral prefrontal cortex (DLPFC), supplemental motor areas (SMA), inferior parietal lobule (IPL), superior parietal lobule (SPL), insula, cerebellum and basal ganglia. A direct comparison revealed that the right DLPFC and the right IPL showed a higher level of activation during the 'Imagery with Ball' than during the 'Imagery'+'Ball' conditions. Our studies suggested that the right front-parietal networks were involved in the motor imagery of an action with an object.

Ipsilateral wrist-ankle movements in the sagittal plane encoded in extrinsic reference frame.

When performing oscillatory movements of two joints in the sagittal plane, there is a directional constraint for performing such movements. Previous studies could not distinguish whether the directional constraint reflected movement direction encoded in the extrinsic (outside the body) reference frame or in the intrinsic (the participants' torso/head) reference frame since participants performed coordinated movements in a sitting position where the torso/head was stationary relative to the external world. In order to discern the reference frame in the present study, participants performed paced oscillatory movements of the ipsilateral wrist and ankle in the sagittal plane in a standing position so that the torso/head moved relative to the external world. The coordinated movements were performed in one of two modes of coordination, moving the hand upward concomitant with either ankle plantarflexion or ankle dorsiflexion. The same directional mode relative to extrinsic space was more stable and accurate as compared with the opposite directional mode. When forearm position was changed from the pronated position to the supinated position, similar results were obtained, indicating that the results were independent of a particular coupling of muscles. These findings suggest that the directional constraint on ipsilateral joints movements in the sagittal plane reflects movement direction encoded in the extrinsic reference frame.

Modulation of corticospinal excitability during acquisition of action sequences by observation.

Excitability of the corticospinal pathway increases during observation of an action. However, how corticospinal excitability changes during observation of sequential actions in the course of acquiring novel skills (observational learning) remains unexplored. To investigate this, we used a previously unpracticed sequence of ten hand postures. Participants were asked to repeat observation and replication of the sequence. This block of observation and replication was repeated 5 times. During observation of a given hand posture (OK sign), motor-evoked potentials (MEPs) elicited by transcranial magnetic stimulation were recorded from hand muscles. In experiment 1, the OK sign appeared in the 9th position of the sequence. Almost all participants could replicate the OK sign only at the 5th block of the experiment. MEP amplitude was greater than that in the control, and decreased with the stages. This suggested that during observational learning of sequential hand postures MEP changed with the progress of the learning. To evaluate this idea, we performed two additional experiments. In experiment 2, the OK sign appeared in the 2nd position. Almost all participants replicated the OK sign even in the 1st block. The MEP amplitude did not change across stages. In experiment 3, the OK sign appeared in the 9th position, but the order of other signs was randomized in every stage. Many participants were not able to replicate the OK sign even during the 5th block of the experiment. The MEP amplitude did not change across stages. These results suggest that: (1) During observational learning modulation of corticospinal excitability is associated with the learning process. (2) Corticospinal excitability decreases as learning progresses.

Influence of somatosensory input on corticospinal excitability during motor imagery.

Our previous studies showed that corticospinal excitability during imagery of squeezing a foam ball was enhanced by somatosensory input generated by passively holding the ball. In the present study, using the same experimental model, we investigated whether corticospinal excitability was influenced by holding the object with the hand opposite to the imagined hand. Corticospinal excitability was assessed by monitoring motor evoked potentials (MEPs) in the first dorsal interosseous muscle following transcranial magnetic stimulation over the motor cortex during motor imagery. Subjects were asked to imagine squeezing a foam ball with the right hand (experiment 1) or the left hand (experiment 2), while either holding nothing (Null condition), a ball in the right hand (Right condition) or a ball in the left hand (Left condition). The MEPs amplitude during motor imagery was increased, only when the holding hand and the imagined hand were on the same side. These results suggest that performance improvement and rehabilitation exercises will be more effective when somatosensory stimulation and motor imagery are done on the same side.

The modulation of corticospinal excitability during motor imagery of actions with objects.

We investigated whether corticospinal excitability during motor imagery of actions (the power or the pincer grip) with objects was influenced by actually touching objects (tactile input) and by the congruency of posture with the imagined action (proprioceptive input). Corticospinal excitability was assessed by monitoring motor evoked potentials (MEPs) in the first dorsal interosseous following transcranial magnetic stimulation over the motor cortex. MEPs were recorded during imagery of the power grip of a larger-sized ball (7 cm) or the pincer grip of a smaller-sized ball (3 cm)--with or without passively holding the larger-sized ball with the holding posture or the smaller-sized ball with the pinching posture. During imagery of the power grip, MEPs amplitude was increased only while the actual posture was the same as the imagined action (the holding posture). On the other hand, during imagery of the pincer grip while touching the ball, MEPs amplitude was enhanced in both postures. To examine the pure effect of touching (tactile input), we recorded MEPs during imagery of the power and pincer grip while touching various areas of an open palm with a flat foam pad. The MEPs amplitude was not affected by the palmer touching. These findings suggest that corticospinal excitability during imagery with an object is modulated by actually touching an object through the combination of tactile and proprioceptive inputs.

Determinant of leg stiffness during hopping is frequency-dependent.

Identifying the major determinant of leg stiffness during hopping would be helpful in the development of more effective training methods. Despite the fact that overall leg stiffness depends on a combination of the joint stiffness, it is unclear how the major determinants of leg stiffness are influenced by hopping frequency. The purpose of this study was to identify the major determinant of leg stiffness over a wide range of hopping frequencies. Fourteen well-trained male athletes performed in a place hopping on two legs, at three frequencies (1.5, 2.2 and 3.0 Hz). We determined leg and joint stiffness of the hip, knee and ankle from kinetic and kinematic data. Multiple linear regression analysis revealed that knee stiffness could explain more of the variance of leg stiffness than could ankle or hip stiffness at 1.5 Hz hopping. Further, only ankle stiffness was significantly correlated with leg stiffness at both 2.2 and 3.0 Hz, and the standardized regression coefficient of ankle stiffness was higher than that of knee and hip stiffness. The results of the present study suggest that the major determinant of leg stiffness during hopping switches from knee stiffness to ankle stiffness when the hopping frequency is increased.

Passive knee movement-induced modulation of the soleus H-reflex and alteration in the fascicle length of the medial gastrocnemius muscle in humans.

In humans, an inhibitory via Ia afferent pathway from the medial gastrocnemius (MG) to the soleus (SOL) motoneuron pool has been suggested. Herein, we examined the relation between MG fascicle length changes and the SOL H-reflex modulation during passive knee movement. Twelve subjects performed static and passive (5 degrees s(-1)) knee movement tasks with the ankle immobilized using an isokinetic dynamometer in sitting posture. The maximal H- and M-waves were measured at four target angles (20 degrees, 40 degrees, 60 degrees, and 80 degrees flexion from full knee extension). The MG fascicles length and velocity were measured using a B-mode ultrasonic apparatus. Results demonstrated that the SOL Hmax/Mmax; i.e., ratio of the maximal H- to M-waves, was attenuated with increasing MG fascicle length in static tasks. The SOL Hmax/Mmax at 20 degrees was significantly attenuated compared with 60 degrees and 80 degrees with increasing MG fascicle length and lengthening velocity in passive knee extension. However, no significant differences in the SOL Hmax/Mmax were found across the target angles in the passive knee flexion task. In conclusion, as muscle spindles increase their discharge with lengthening fascicle velocity, but keep silent when fascicles shorten, our data suggest that lengthening the MG facilitates an inhibitory Ia pathway from MG to SOL, and modulates SOL motoneuron activity during movements.