Temporal aspects of passive movement-related corticomotor inhibition

We have previously shown that during rhythmic passive movement of the index finger, the amplitude of the motor evoke potential (MEP) of the first dorsal interosseous muscle (FDI) as the index finger moved through mid-range adduction, is significantly reduced compared to rest [Edwards, D. J., Thickbroom, G. W., Byrnes, M. L., Ghosh, S., & Mastaglia, F. L. (2002). Reduced corticomotor excitability with passive movement: A study using Transcranial Magnetic Stimulation. Human Movement Science 21, 533-540]. In the present study we have investigated the time-course of this phenomenon. We found that MEP amplitude was significantly reduced at the mid-range position in the first cycle of movement (50+/-6% of resting baseline values), and did not vary across subsequent cycles (10 cycles in 50 s), but that MEP amplitude returned to baseline values within 1s of cessation of movement. The results suggest that the pattern of afferent discharge set up by the kinematics of the movement acting at spinal or supraspinal levels underlies the inhibition observed, rather than an effect of central origin or a cumulative effect of ongoing cyclic movement.     

Rapid plastic changes of human primary motor cortex with repetitive motor practice and transcranial magnetic stimulation

Excitability changes of human primary motor cortex are assumed to be associated with motor learning processes. To examine motor behavioral and neural mechanisms in these processes, the adaptive motor learning processes of the index finger abduction were investigated using motor evoked potential (MEP) elicited from the first dorsal interosseous and extensor carpi radialis muscles. Practice effects were examined on changes of MEP amplitudes elicited from these muscles during motor imagery. Given general consensus that the MEP amplitude change during motor imagery is a useful parameter reflecting changes in excitability of the human primary motor cortex, the present results, that MEP amplitudes of both muscles increased with repeated practice by the index finger abduction and that magnitudes of MEP amplitudes of both muscles (motor learning curves) were clearly different, suggested that participation of the muscles performing the index finger abduction gradually changed with practice. Short-term plastic changes of human primary motor cortex occur with repetitive practice and such adaptive change in human primary motor cortex is expressed in human voluntary movement that becomes more automatic.     

Premovement facilitation of corticospinal excitability before simple and sequential movement

The purpose of this study was to investigate whether premovement facilitation of corticospinal excitability before sequential movement was different from that before simple movement. Each of 7 participants who performed choice reaction tasks with the right hand pressed a force transducer with the index finger in response to a start cue or pressed the transducers sequentially with the index finger, little finger, thumb, little finger, and index finger. Transcranial magnetic stimulation was delivered to the left motor cortex before the electromyographic burst in the first dorsal interosseous muscle and motor evoked potentials were recorded from the first dorsal interosseous muscle. The amplitude of the motor-evoked potential increased as its onset got closer to the onset of the electromyographic burst. The increase before the sequential movement was larger and began earlier than that before the simple movement. These findings indicate that premovement facilitation of corticospinal excitability is different in magnitude and timing between sequential and simple movements.     

Having a body versus moving your body: How agency structures body-ownership

We investigated how motor agency in the voluntary control of body movement influences body awareness. In the Rubber Hand Illusion (RHI), synchronous tactile stimulation of a rubber hand and the participant's hand leads to a feeling of the rubber hand being incorporated in the participant's own body. One quantifiable behavioural correlate of the illusion is an induced shift in the perceived location of the participant's hand towards the rubber hand. Previous studies showed that the induced changes in body awareness are local and fragmented: the proprioceptive drift is largely restricted to the stimulated finger. In the present study, we investigated whether active and passive movements, rather than tactile stimulation, would lead to similarly fragmented body awareness. Participants watched a projected image of their hand under three conditions: active finger movement, passive finger movement, and tactile stimulation. Visual feedback was either synchronous or asynchronous with respect to stimulation of the hand. A significant overall RHI, defined as greater drifts following synchronous than asynchronous stimulation, was found in all cases. However, the distribution of the RHI across stimulated and non-stimulated fingers depended on the kind of stimulation. Localised proprioceptive drifts, specific to the stimulated finger, were found for tactile and passive stimulation. Conversely, during active movement of a single digit, the proprioceptive drifts were not localised to that digit, but were spread across the whole hand. Whereas a purely proprioceptive sense of body-ownership is local and fragmented, the motor sense of agency integrates distinct body-parts into a coherent, unified awareness of the body.     

Intrasensory attention: kinaesthetic versus cutaneous inputs

Blindfolded participants felt pairs of raised-line drawings simultaneously, one with each index finger. The stimuli presented at each fingertip were 180 degrees rotations of each other (eg 6 and 9). One finger moved (either actively or passively), and this in turn caused movement of a matched raised line underneath the stationary finger on the other hand, in a yoked manner. Thus, a 6 at the moving finger would be felt as a 9 on the stationary finger. On all trials there was a raised line moving underneath the stationary fully passive finger. For the moving finger, a raised line was present on only half of the trials. When a raised line could be felt at the moving fingertip, the shape followed by this finger was more often reported than was the shape present at the other (stationary) fingertip. However, when no line was present under the moving finger (ie when movement became the major cue for shape), subjects reported experiencing the shape moved under the stationary fingertip. Results are interpreted as an indication that cutaneous information can be more 'attention-getting' than kinaesthetic information, and are considered to support the modality-appropriateness theory.     

Effects of feedback from active and passive body parts on spatial and temporal parameters in sensorimotor synchronization

Previous research on sensorimotor synchronization has manipulated the somatosensory information received from the tapping finger to investigate how feedback from an active effector affects temporal coordination. The current study explored the role of feedback from passive body parts in the regulation of spatiotemporal motor control parameters by employing a task that required finger tapping on one’s own skin at anatomical locations of varying tactile sensitivity. A motion capture system recorded participants’ movements as they synchronized with an auditory pacing signal by tapping with the right index finger on either their left index fingertip (Finger/Finger) or forearm (Finger/Forearm). Results indicated that tap timing was more variable, and movement amplitude was larger and more variable, when tapping on the finger than when tapping on the less sensitive forearm. Finger/Finger tapping may be impaired relative to Finger/Forearm tapping due to ambiguity arising through overlap in neural activity associated with tactile feedback from the active and the passive limb in the former. To compensate, the control system may strengthen the assignment of tap-related feedback to the active finger by generating correlated noise in movement kinematics and tap dynamics.     

The Effect of Paired Muscle Stimulation on Preparation for Movement

Paired muscle stimulation is used clinically to facilitate the performance of motor tasks for individuals with motor dysfunction. However, the optimal temporal relationship between stimuli for enhancing movement remains unknown. We hypothesized that synchronous, muscle stimulation would increase the extent to which stimulated muscles are concurrently prepared for movement. We validated a measure of muscle-specific changes in corticomotor excitability prior to movement. We used this measure to examine the preparation of the first dorsal interosseous (FDI), abductor digiti minimi (ADM), abductor pollicis brevis (APB) muscles prior to voluntary muscle contractions before and after paired muscle stimulation at four interstimulus intervals (0, 5, 10, and 75 ms). Paired muscle stimulation increased premovement excitability in the stimulated FDI, but not in the ADM muscle. Interstimulus interval was not a significant factor in determining efficacy of the protocol. Paired stimulation, therefore, did not result in a functional association being formed between the stimulated muscles. Somatosensory potentials evoked by the muscle stimuli were small compared to those commonly elicited by stimulation of peripheral nerves, suggesting that the lack of functional association formation between muscles may be due to the small magnitude of afferent volleys from the stimulated muscles, particularly the ADM, reaching the cortex.     

Shifts in kinesthesis through time and after active and passive movement.

The position sense of a stationary arm was investigated subsequent to an horizontally adductive movement with axis the shoulder joint. The right arm was the treated arm: it reached a test position actively, using minimal voluntary effort, or passively from each of 10 starting positons. The blind-folded S localized the index finger of the treated arm by attempting to touch it with the index finger of his left hand. The results indicate that subsequent to active movement the final position of a limb is more accurately known than a position resulting from passive movement. A second finding is that concomitant with both forms of limb placement there is a unidirectional drift of perceived limb position over trials.     

Time and direction preparation of the long latency stretch reflex

This study investigated time and direction preparation of motor response to force load while intending to maintain the finger at the initial neutral position. Force load extending or flexing the index finger was given while healthy humans intended to maintain the index finger at the initial neutral position. Electromyographic activity was recorded from the first dorsal interosseous muscle. A precue with or without advanced information regarding the direction of the forthcoming force load was given 1000 ms before force load. Trials without the precue were inserted between the precued trials. A long latency stretch reflex was elicited by force load regardless of its direction, indicating that the long latency stretch reflex is elicited not only by muscle stretch afferents, but also by direction-insensitive sensations. Time preparation of motor response to either direction of force load enhanced the long latency stretch reflex, indicating that time preparation is not mediated by afferent discharge of muscle stretch. Direction preparation enhanced the long latency stretch reflex and increased corticospinal excitability 0–20 ms after force load when force load was given in the direction stretching the muscle. These enhancements must be induced by preset of the afferent pathway mediating segmental stretch reflex.     

Aspects of body self-calibration

The representation of body orientation and configuration is dependent on multiple sources of afferent and efferent information about ongoing and intended patterns of movement and posture. Under normal terrestrial conditions, we feel virtually weightless and we do not perceive the actual forces associated with movement and support of our body. It is during exposure to unusual forces and patterns of sensory feedback during locomotion that computations and mechanisms underlying the ongoing calibration of our body dimensions and movements are revealed. This review discusses the normal mechanisms of our position sense and calibration of our kinaesthetic, visual and auditory sensory systems, and then explores the adaptations that take place to transient Coriolis forces generated during passive body rotation. The latter are very rapid adaptations that allow body movements to become accurate again, even in the absence of visual feedback. Muscle spindle activity interpreted in relation to motor commands and internally modeled reafference is an important component in permitting this adaptation. During voluntary rotary movements of the body, the central nervous system automatically compensates for the Coriolis forces generated by limb movements. This allows accurate control to be maintained without our perceiving the forces generated.     

Observing how others lift light or heavy objects: time-dependent encoding of grip force in the primary motor cortex

During movement observation, corticomotor excitability of the observer's primary motor cortex (M1) is modulated according to the force requirements of the observed action. Here, we explored the time course of observation-induced force encoding. Force-related changes in M1-excitability were assessed by delivering transcranial magnetic stimulations at distinct temporal phases of an observed reach-grasp-lift action. Temporal changes in force-related electromyographic activity were also assessed during active movement execution. In observation conditions in which a heavy object was lifted, M1-excitability was higher compared to conditions in which a light object was lifted. Both during observation and execution, differential force encoding tended to gradually increase from the grasping phase until the late lift phase. Surprisingly, however, during observation, force encoding was already present at the early reach phase: a time point at which no visual cues on the object's weight were available to the observer. As the observer was aware that the same weight condition was presented repeatedly, this finding may indicate that prior predictions concerning the upcoming weight condition are reflected by M1 excitability. Overall, findings may provide indications that the observer's motor system represents motor predictions as well as muscular requirements to infer the observed movement goal.     

The effects of voluntary movements on auditory-haptic and haptic-haptic temporal order judgments

In two experiments we investigated the effects of voluntary movements on temporal haptic perception. Measures of sensitivity (JND) and temporal alignment (PSS) were obtained from temporal order judgments made on intermodal auditory-haptic (Experiment 1) or intramodal haptic (Experiment 2) stimulus pairs under three movement conditions. In the baseline, static condition, the arm of the participants remained stationary. In the passive condition, the arm was displaced by a servo-controlled motorized device. In the active condition, the participants moved voluntarily. The auditory stimulus was a short, 500Hz tone presented over headphones and the haptic stimulus was a brief suprathreshold force pulse applied to the tip of the index finger orthogonally to the finger movement. Active movement did not significantly affect discrimination sensitivity on the auditory-haptic stimulus pairs, whereas it significantly improved sensitivity in the case of the haptic stimulus pair, demonstrating a key role for motor command information in temporal sensitivity in the haptic system. Points of subjective simultaneity were by-and-large coincident with physical simultaneity, with one striking exception in the passive condition with the auditory-haptic stimulus pair. In the latter case, the haptic stimulus had to be presented 45ms before the auditory stimulus in order to obtain subjective simultaneity. A model is proposed to explain the discrimination performance.     

Anomalous control: when 'free-will' is not conscious

The conscious feeling of exercising 'free-will' is fundamental to our sense of self. However, in some psychopathological conditions actions may be experienced as involuntary or unwilled. We have used suggestion in hypnosis to create the experience of involuntariness (anomalous control) in normal participants. We compared a voluntary finger movement, a passive movement and a voluntary movement suggested by hypnosis to be 'involuntary.' Hypnosis itself had no effect on the subjective experience of voluntariness associated with willed movements and passive movements or on time estimations of their occurrence. However, subjective time estimates of a hypnotically-suggested, 'involuntary' finger movement were more similar to those for passive movements than for voluntary movements. The experience of anomalous control is qualitatively and quantitatively different from the normal conscious experience of a similar act produced intentionally. The experience of anomalous control may be produced either by pathology, or, in our case, by suggestion.     

Modulations of use-dependent excitability changes of human motor cortex (M1) by practice condition

Effects of repetitive index finger abductions on excitability changes in the human primary motor cortex (Ml) are assumed to be dependent on practice conditions of the task. To address how different effects occur dependent on various practice conditions, motor evoked potentials (MEPs) elicited from the first dorsal interosseous (FDI) muscle were investigated. Practice effects on the index finger abduction were examined for changes in excitability of first dorsal interosseous muscle under three forearm position changes (neutral vs prone) and two muscle contraction modes (isometric vs isotonic). Analysis showed that after practice MEP amplitude increased in the prone position but not in the neutral position and MEP increases in the isotonic contraction were larger than those in the isometric mode. These results suggest that use-dependent excitability changes are largely dependent on practice conditions because the amount of afferent input depends on the practice conditions.     

Proprioceptive feedback as a mediator in interlimb timing

Prior findings regarded as evidence for proprioceptive feedback as a mediator in interlimb timing can also be interpreted as evidence for motor outflow because they came from research that had subjects make voluntary movements, and such movements allow for both feedback and outflow to operate. The present study was designed to resolve this controversy by determining if these findings could be replicated with passive movements which allow for feedback, but not outflow, to operate. The interlimb timing task studied was one where subjects made the timing response with their right hand while moving their left arm during the 1.5-sec interval to be timed. Three groups of 16 male college students performed 50 trials of the right-hand response with knowledge of results, under one of three left-arm conditions: (a) passive movement, (b) voluntary movement, and (c) no movement. The results indicated that the findings were replicated with passive movements and this was interpreted as support for the involvement of proprioceptive feedback in interlimb timing.     

Emotional state and initiating cue alter central and peripheral motor processes

Evidence indicates that voluntary and involuntary movements are altered by affective context as well as the characteristics of an initiating cue. The purpose of this study was to determine the contribution of central and peripheral mechanisms to this phenomenon. During the presentation of pleasant, unpleasant, neutral, and blank images, participants (N = 33) responded to auditory stimuli (startle, 107 dB startle or 80 dB tone) by initiating a bimanual isometric contraction of the wrist and finger extensor muscles. Analyses of electromyography and force measures supported the hypothesis that exposure to unpleasant images accelerates central processing times and increases the gradient of slope of peripheral movement execution. In addition, startle cues as compared with tone cues accelerated and magnified all temporal and amplitude indices. Collectively, these findings have noteworthy implications for (a) those seeking to facilitate the speed and force of voluntary movement (i.e., movement rehabilitation), (b) understanding the higher incidence of motor difficulty in individuals with affective disorders, and (c) those seeking to regulate emotional input so as to optimize the quality of intended movements.     

Musical groove modulates motor cortex excitability: A TMS investigation

Groove is often described as a musical quality that can induce movement in a listener. This study examines the effects of listening to groove music on corticospinal excitability. Musicians and non-musicians listened to high-groove music, low-groove music, and spectrally matched noise, while receiving single-pulse transcranial magnetic stimulation (TMS) over the primary motor cortex either on-beat or off-beat. We examined changes in the amplitude of the motor-evoked potentials (MEPs), recorded from hand and arm muscles, as an index of activity within the motor system. Musicians and non-musicians rated groove similarly. MEP results showed that high-groove music modulated corticospinal excitability, whereas no difference occurred between low-groove music and noise. More specifically, musicians’ MEPs were larger with high-groove than low-groove music, and this effect was especially pronounced for on-beat compared to off-beat pulses. These results indicate that high-groove music increasingly engages the motor system, and the temporal modulation of corticospinal excitability with the beat could stem from tight auditory–motor links in musicians. Conversely, non-musicians’ MEPs were smaller for high-groove than low-groove music, and there was no effect of on- versus off-beat pulses, potentially stemming from suppression of overt movement. In sum, high-groove music engages the motor system, and previous training modulates how listening to music with a strong groove activates the motor system.     

Busy doing nothing: Evidence for nonaction-effect binding

Research on voluntary action has focused on the question of how we represent our behavior on a motor and cognitive level. However, the question of how we represent voluntary not acting has been completely neglected. The aim of the present study was to investigate the cognitive and motor representation of intentionally not acting. By using an action-effect binding approach, we demonstrate similarities of action and nonaction. In particular, our results reveal that voluntary nonactions can be bound to an effect tone. This finding suggests that effect binding is not restricted to an association between a motor representation and a successive effect (action-effect binding) but can also occur for an intended nonaction and its effect (nonaction-effect binding). Moreover, we demonstrate that nonactions have to be initiated voluntarily in order to elicit nonaction-effect binding.     

Effects of proprioceptive neuromuscular facilitation on the initiation of voluntary movement and motor evoked potentials in upper limb muscles

To better understand the mechanisms behind proprioceptive neuromuscular facilitation (PNF), an important method in motor rehabilitation, we investigated the effects of assuming a PNF posture relative to a neutral posture on the initiation of voluntary movement (Experiment 1) and the excitability of the motor cortex (Experiment 2) using a wrist extension task. The initiation of voluntary wrist movement was operationalized in terms of the electromyographic reaction time (EMG-RT), and the excitability of the motor cortex in terms of motor evoked potentials (MEPs). Compared to the neutral position, we found that (1) the facilitation position changed the muscle discharge order enhancing the movement efficiency of the joint, (2) the facilitation position led to a reduction in EMG-RT, the magnitude of which depended on the proximity of the muscle to the movement joint, and (3) MEP amplitude increased and MEP latency decreased in the facilitation position as a function of the proximity of the muscle to the joint. These findings corroborate the presumed effects of PNF and provide insights into the neurophysiological mechanisms underlying the PNF method.     

Finger inter-dependence: linking the kinetic and kinematic variables

We studied the dependence between voluntary motion of a finger and pressing forces produced by the tips of other fingers of the hand. Participants moved one of the fingers (task finger) of the right hand trying to follow a cyclic, ramp-like flexion-extension template at different frequencies. The other fingers (slave fingers) were restricted from moving; their flexion forces were recorded and analyzed. Index finger motion caused the smallest force production by the slave fingers. Larger forces were produced by the neighbors of the task finger; these forces showed strong modulation over the range of motion of the task finger. The enslaved forces were higher during the flexion phase of the movement cycle as compared to the extension phase. The index of enslaving expressed in N/rad was higher when the task finger moved through the more flexed postures. The dependence of enslaving on both range and direction of task finger motion poses problems for methods of analysis of finger coordination based on an assumption of universal matrices of finger interdependence.