Visual tracking and entrainment to an environmental rhythm

This study investigated the role that visual tracking plays in coupling rhythmic limb movements to an environmental rhythm. Two experiments were conducted in which participants swung a hand-held pendulum while tracking an oscillating stimulus or while keeping their eyes fixed on a stationary location directly above an oscillating stimulus. It was expected that the participants' rhythmic movements would become entrained to the oscillating stimulus in both conditions but that visual tracking would strengthen this entrainment. Experiment 1 investigated the role of visual tracking in establishing unintentional entrainment. Experiment 2 investigated the role of visual tracking in intentional entrainment. As predicted, participants exhibited greater unintentional coordination and more stable intentional coordination when they tracked the stimulus. These findings highlight the importance of understanding the role of eye movements in environmental coordination.     

Informational constraints on spontaneous visuomotor entrainment

Past research has revealed that an individual’s rhythmic limb movements become spontaneously entrained to an environmental rhythm if visual information about the rhythm is available and its frequency is near that of the individual’s movements. Research has also demonstrated that if the eyes track an environmental stimulus, the spontaneous entrainment to the rhythm is strengthened. One hypothesis explaining this enhancement of spontaneous entrainment is that the limb movements and eye movements are linked through a neuromuscular coupling or synergy. Another is that eye-tracking facilitates the pick up of important coordinating information. Experiment 1 investigated the first hypothesis by evaluating whether any rhythmic movement of the eyes would facilitate spontaneous entrainment. Experiments 2 and 3 (respectively) explored whether eye-tracking strengthens spontaneous entrainment by allowing the pickup of trajectory direction change information or allowing an increase in the amount of information to be picked-up. Results suggest that the eye-tracking enhancement of spontaneous entrainment is a consequence of increasing the amount of information available to be picked-up.     

Dependence of Finger Flexion Force on the Posture of the Nonperforming Fingers During Key Pressing Tasks

In visuomotor tasks that involve accuracy demands, small directional changes in the trajectories have been taken as evidence of feedback-based error corrections. In the present study variability, or intermittency, in visuomanual tracking of sinusoidal targets was investigated. Two lines of analyses were pursued: First, the hypothesis that humans fundamentally act as intermittent servo-controllers was re-examined, probing the question of whether discontinuities in the movement trajectory directly imply intermittent control. Second, an alternative hypothesis was evaluated: that rhythmic tracking movements are generated by entrainment between the oscillations of the target and the actor, such that intermittency expresses the degree of stability. In 2 experiments, participants (N = 6 in each experiment) swung 1 of 2 different hand-held pendulums, tracking a rhythmic target that oscillated at different frequencies with a constant amplitude. In 1 line of analyses, the authors tested the intermittency hypothesis by using the typical kinematic error measures and spectral analysis. In a 2nd line, they examined relative phase and its variability, following analyses of rhythmic interlimb coordination. The results showed that visually guided corrective processes play a role, especially for slow movements. Intermittency, assessed as frequency and power components of the movement trajectory, was found to change as a function of both target frequency and the manipulandum's inertia. Support for entrainment was found in conditions in which task frequency was identical to or higher than the effector's eigenfrequency. The results suggest that it is the symmetry between task and effector that determines which behavioral regime is dominant.     

Period basin of entrainment for unintentional visual coordination

Researchers have demonstrated that a person's rhythmic movements can become unintentionally entrained to another person's rhythmic movements or an environmental event. There are indications, however, that in both cases the likelihood of entrainment depends on the difference between the uncoupled periods of the two rhythms. The authors examined the range of period differences over which unintentional visual coordination might occur in 16 participants (Experiment 1) and 15 participants (Experiment 2). Cross-spectral coherence analysis and the distribution of continuous relative phase revealed that visual entrainment decreased as the difference between participants' preferred period and the experimenter-determined period of the environmental stimulus increased. The present findings extend the dynamical systems perspective on person-environment coupling and highlight the significance of period difference to the emergence of unintentional coordination.     

Rapid motor adaptations to subliminal frequency shifts during syncopated rhythmic sensorimotor synchronization

The synchronization of rhythmic arm movements to a syncopated metronome cue was studied in a step-change design whereby small tempo shifts were inserted at fixed time points into the metronome frequency. The cueing sequence involved three stimulus types: (1) target contact in synchrony with the metronome beats, (2) syncopated target contact midway in time between audible beats, and (3) syncopated target contact following either a +2% or -2% change in stimulus frequency. Analysis of normalized and aggregated data revealed that (1) during the syncopation condition the response period showed a rapid adaptation to the frequency-incremented stimulus period, (2) response period was less variable during syncopated movement, (3) mean synchronization error and variability, calculated during syncopation relative to the mathematical midpoint of the stimulus cycle, were reduced during syncopated movements, and (4) synchronization error following the frequency increment showed trends to return linearly to pre-increment values which was fully achieved in the -2% change condition only. The results suggest that frequency entrainment to stimulus period was possible during syncopated movement with the response and stimulus onsets 180 degrees out of phase. Most remarkably, 70-80% of the adaptation of the response period to the new stimulus period was immediately attained during the second half cycle of the syncopated movement. Finally, a mathematical model, based on recursion, was introduced that accurately modeled actual data as a function of the previous stimulus and response intervals and a weighted response of period error and synchronization error, which showed dominance of frequency entrainment over phase entrainment during rhythmic synchronization.     

Synergistic influences of sensory and central stimuli on non-voluntary rhythmic arm movements

In recent years, neuromodulation of the cervical spinal circuitry has become an area of interest for investigating rhythmogenesis of the human spinal cord and interaction between cervical and lumbosacral circuitries, given the involvement of rhythmic arm muscle activity in many locomotor tasks. We have previously shown that arm muscle vibrostimulation can elicit non-voluntary upper limb oscillations in unloading body conditions. Here we investigated the excitability of the cervical spinal circuitry by applying different peripheral and central stimuli in healthy humans. The rationale for applying combined stimuli is that the efficiency of only one stimulus is generally limited. We found that low-intensity electrical stimulation of the superficial arm median nerve can evoke rhythmic arm movements. Furthermore, the movements were enhanced by additional peripheral stimuli (e.g., arm muscle vibration, head turns or passive rhythmic leg movements). Finally, low-frequency transcranial magnetic stimulation of the motor cortex significantly facilitated rhythmogenesis. The findings are discussed in the general framework of a brain-spinal interface for developing adaptive central pattern generator-modulating therapies.     

Contours of equal perceived amplitude and equal perceived frequency for electrocutaneous stimuli

Previous measurements of equal-sensation contours for electrocutaneous stimuli consisting of repeated bursts of biphasic pulses have shown that stimulus frequency has little effect on perceived amplitude, and that stimulus amplitude has no effect on perceived frequency. These earlier contours, however, were measured over a very restricted range of amplitude and frequency or for a single perceived amplitude or perceived frequency. Contours of equal perceived amplitude and equal perceived frequency were measured in the present study for stimuli covering most of the useable range of amplitudes and frequencies: 3–12 dB SL and 4–64 Hz. Eight naive subjects generated contours of equal perceived amplitude at four reference amplitudes via Békésy tracking, and 8 additional subjects generated contours of equal perceived frequency at three reference frequencies. The contours of equal perceived amplitude declined slightly but significantly with increases in stimulus frequency, consistent with previous results. The shape of the contours was also slightly dependent on the amplitude of the reference stimulus. Contours of equal perceived frequency were unaffected by stimulus amplitude on the average, but the contour shape did vary modestly, though erratically, with reference frequency.     

Effects of visual and verbal interaction on unintentional interpersonal coordination

Previous research has demonstrated that people's movements can become unintentionally coordinated during interpersonal interaction. The current study sought to uncover the degree to which visual and verbal (conversation) interaction constrains and organizes the rhythmic limb movements of coactors. Two experiments were conducted in which pairs of participants completed an interpersonal puzzle task while swinging handheld pendulums with instructions that minimized intentional coordination but facilitated either visual or verbal interaction. Cross-spectral analysis revealed a higher degree of coordination for conditions in which the pairs were visually coupled. In contrast, verbal interaction alone was not found to provide a sufficient medium for unintentional coordination to occur, nor did it enhance the unintentional coordination that emerged during visual interaction. The results raise questions concerning differences between visual and verbal informational linkages during interaction and how these differences may affect interpersonal movement production and its coordination.     

Limb Segment Recruitment as Function of Movement Direction,Amplitude, and Speed

Coordination of limb segments in graphic motor behavior has been studied primarily in cyclic tasks. In the present study, limb segment recruitment patterns were investigated in a discrete line-drawing task. Subjects (N = 11) performed pointing movements varying in direction, amplitude, and speed. The contributions of index finger, hand, and arm to the movement were analyzed by evaluating the angular displacements in 7 joint dimensions. The results showed that amplitude and direction affected limb segment involvement in the same way they have been reported to affect it in cyclic movements. Upward left- (up-left) directed movements were primarily achieved by fingers and arm, whereas upward right- (up-right) directed movements were accomplished with the hand and the arm. Large amplitudes elicited not only an increase of proximal but also a decrease of distal limb segment involvement, especially in the up-left direction. In the present discrete pointing task, effects of speed on limb segment involvement were different from speed effects that were observed earlier in cyclic tasks: Larger limb segments became more involved in fast than in slow discrete movements. With respect to the timing of limb segment recruitment, all joints tended to move simultaneously, but small deviations from synchronous joint movement onset and offset were present. The results are discussed in the context of recent theories of limb segment coordination.     

Auditory cueing facilitates temporospatial accuracy of sequential movements

Effectively executing goal-directed behaviours requires both temporal and spatial accuracy. Previous work has shown that providing auditory cues enhances the timing of upper-limb movements. Interestingly, alternate work has shown beneficial effects of multisensory cueing (i.e., combined audiovisual) on temporospatial motor control. As a result, it is not clear whether adding visual to auditory cues can enhance the temporospatial control of sequential upper-limb movements specifically. The present study utilized a sequential pointing task to investigate the effects of auditory, visual, and audiovisual cueing on temporospatial errors. Eighteen participants performed pointing movements to five targets representing short, intermediate, and large movement amplitudes. Five isochronous auditory, visual, or audiovisual priming cues were provided to specify an equal movement duration for all amplitudes prior to movement onset. Movement time errors were then computed as the difference between actual and predicted movement times specified by the sensory cues, yielding delta movement time errors (ΔMTE). It was hypothesized that auditory-based (i.e., auditory and audiovisual) cueing would yield lower movement time errors compared to visual cueing. The results showed that providing auditory relative to visual priming cues alone reduced ΔMTE particularly for intermediate amplitude movements. The results further highlighted the beneficial impact of unimodal auditory cueing for improving visuomotor control in the absence of significant effects for the multisensory audiovisual condition.     

Electromyographic activity,H-reflex modulation and corticospinal input to forearm motoneurones during active and passive rhythmic movements

Four subjects produced coordinated movements, consisting of flexion and extension of the wrist in ipsilateral (right wrist only), contralateral (left wrist only), inphase (both wrists in flexion or both in extension) and antiphase (one wrist in flexion, the other in extension) conditions. Electromyographic (EMG) activity was recorded from right wrist flexor and extensor muscles. In one session, transcranial magnetic stimuli (TMS) of the left motor cortex, around threshold intensity, evoked short-latency responses in the right wrist extensors and flexors. In another session, the median nerve at the cubital fossa was stimulated to elicit an H-reflex in the right flexor carpi radialis (rFCR). A movement cycle was divided into 8 segments. In total, 10 identical stimuli were delivered during each segment in each condition, at two movement frequencies. The magnitude of the EMG reponses to TMS was modulated markedly during movements made in the ipsilateral condition, and in both bimanual conditions. EMG activity was greater, and motor-evoked potentials (MEPs) were larger in the antiphase condition than in the inphase condition. When the amplitudes of the MEPs were normalised with respect to background EMG, no significant differences between the bimanual conditions were obtained. For H-reflexes, significant differences between the two bimanual conditions were observed, suggesting differences in levels of excitability of the Ia afferent pathway. These differences were attributed to segmental input associated with changes in muscle length arising from limb movement, and upon descending input to the spinal cord, possibly mediated by Renshaw cell inhibition. During rhythmic passive movement of the right limb, H-reflexes were inhibited and MEPs potentiated in a cyclic fashion. Passive movement of the contralateral left limb resulted in inhibition of both responses.PsycINFO classification: 2330; 2530; 2540     

Oculomanual Coordination in Tracking of Pseudorandom Target Motion Stimuli

Oculomanual coordination was investigated in 9 healthy subjects during tracking of pseudorandom motion stimuli. Each subject was required to track visual stimuli under eye-hand (EH) and eye-alone (EA) conditions. Subjects were exposed to 3 types of mixed sinusoidal stimulus with varying frequency or amplitude of the highest frequency component, or various degrees of irregularity. Progressive degradation in tracking performance was nonlinearly induced by an increase in either (a) the highest frequency component or (b) its amplitude, but not by stimulus irregularity. No significant difference was found in eye velocity gain and phase under the EH and EA conditions. Eye and hand responses were found to be highly correlated in gain and phase when compared across frequencies and motion stimuli. The results suggest that frequency and amplitude are dominant factors controlling the breakdown of oculomanual performance in response to pseudorandom stimuli. Frequency responses of smooth pursuit eye movements are not affected by the hand motion in pursuit of unpredictable stimuli. Eye and hand motor systems appear to share common nonlinear drive mechanisms when pursuing pseudorandom target motion stimuli.     

Backward and forward masking associated with saccadic eye movement

Visual masking effects on test flash thresholds were measured under real and simulated eye movement conditions to determine whether visual masking is primarily responsible for elevations in threshold that are sometimes associated with saccadic eye movements. Brief luminous flashes presented to the central retina before, during, and after saccades were masked by stimuli presented either pre- or postsaccadically. The amount and time course of masking were quantitatively dependent on stimulus parameters of intensity and temporal separation and were unaffected by eye movement parameters (amplitude, velocity, direction) as long as retinal stimulus conditions were constant. The duration of forward masking was longer than that of backward masking. When retinal conditions during saccades were mimicked while the eyes were held steady, masking interactions were identical to those obtained during real saccades. These results indicate that masking effects during saccades in ordinary environments are determined solely by the stimulus situation at the retina. Putative nonvisual, centrally originating saccadic suppression suggested by other authors is evidently not additive with visually determined masking during saccades.     

The effect of mechanical context on attentional cost in unimanual coordination

The purpose of this study was to address the attentional cost of sensorimotor coordination by determining if changes to the mechanical context of movement would influence the ability to attend and respond to an alternate stimulus. Nine right-handed participants performed rhythmic pronation and supination movements of the forearm in time with an auditory metronome. A secondary task, consisting of a pedal response to visual probe stimuli, was employed to infer the attentional cost of the coordination task. When the axis-of-rotation (AOR) was placed below the long axis of the forearm, the average time to react (RT) to the probe stimuli was greater for the supinate-on-the-beat condition than for the pronate-on-the-beat condition. Conversely, with the AOR above the forearm, RT for the pronate-on-the-beat pattern was greater than that for the supinate-on-the-beat pattern. Thus, changing the mechanical context of an upper limb coordination task altered the central processing cost required to maintain pattern stability. This finding provides further evidence that the attentional resources required to produce a particular movement are determined by the ease with which the action is executed by the sensorimotor system.     

Interference effects in bimanual coordination are independent of movement type

Simultaneously executed limb movements interfere with each other. Whereas the interference between discrete movements is examined mostly from a cognitive perspective, that between rhythmic movements is studied mainly from a dynamical systems perspective. As the tools and concepts developed by both communities are limited in their applicability to the other domain, it remains unclear if a common cause underlies motor interference in both domains. We investigated the interference between simultaneously executed discrete and rhythmic wrist movements. The discrete movements' reaction time and movement time decreased with increasing rhythmic movement frequency. The discrete movements accelerated or decelerated the rhythmic movements in a manner that depended on movement frequency and the discrete movement's initiation phase. The acceleration/deceleration profile was bimodal at low frequencies and unimodal at high frequencies, mimicking the hallmark feature of rhythmic-rhythmic coordination, thus suggesting that interference between movements may be invariant across different movement types.     

The role of the adjacency between background cues and objects in visual localization during ocular pursuit

Subjects used eye movements to pursue a light target that moved from left to right with a velocity of 15 deg s-1. The stimulus was a sudden five-fold decrease in target intensity during the movement. The subject's task was to localize the stimulus relative to either a single stationary background point or the midpoint between two points (28 deg apart) placed 0.5 deg above the target path. The stimulus was usually mislocated in the direction of eye movement; the mislocation was affected by the spatial adjacency between background and stimulus. When an auditory, rather than a visual, stimulus was presented during tracking, target position at the time of stimulus presentation was visually mislocated in the direction opposite to that of eye movement. The effect of adjacency between background and target remained the same. The involvement of processes of subject-relative and object-relative visual perception is discussed.     

Effects of motor intention on the perception of somatosensory events: A behavioural and functional magnetic resonance imaging study

The intention to execute a movement can modulate our perception of sensory events, and this modulation is observed ahead of both ocular and upper limb movements. However, theoretical accounts of these effects, and also the empirical data, are often contradictory. Accounts of “active touch”, and the premotor theory of attention, have emphasized how movement intention leads to enhanced perceptual processing at the target of a movement, or on the to-be-moved effector. By contrast, recent theories of motor control emphasize how internal “forward” model (FM) estimates may be used to cancel or attenuate sensory signals that arise as a result of self-generated movements. We used behavioural and functional brain imaging (functional magnetic resonance imaging, fMRI) to investigate how perception of a somatosensory stimulus differed according to whether it was delivered to a hand that was about to execute a reaching movement or the alternative, nonmoving, hand. The results of our study demonstrate that a somatosensory stimulus delivered to a hand that is being prepared for movement is perceived to have occurred later than when that same stimulus is delivered to a nonmoving hand. This result indicates that it takes longer for a tactile stimulus to be detected when it is delivered to a moving limb and may correspond to a change in perceptual threshold. Our behavioural results are paralleled by the results of our fMRI study that demonstrated that there were significantly reduced blood-oxygen-level-dependent (BOLD) responses within the parietal operculum and insula following somatosensory stimulation of the hand being prepared for movement, compared to when an identical stimulus was delivered to a nonmoving hand. These findings are consistent with the prediction of FM accounts of motor control that postulate that central sensory suppression of somatosensation accompanies self-generated limb movements, and with previous reports indicating that effects of sensory suppression are observed in higher order somatosensory regions.     

Effects of motor intention on the perception of somatosensory events: a behavioural and functional magnetic resonance imaging study

The intention to execute a movement can modulate our perception of sensory events, and this modulation is observed ahead of both ocular and upper limb movements. However, theoretical accounts of these effects, and also the empirical data, are often contradictory. Accounts of "active touch", and the premotor theory of attention, have emphasized how movement intention leads to enhanced perceptual processing at the target of a movement, or on the to-be-moved effector. By contrast, recent theories of motor control emphasize how internal "forward" model (FM) estimates may be used to cancel or attenuate sensory signals that arise as a result of self-generated movements. We used behavioural and functional brain imaging (functional magnetic resonance imaging, fMRI) to investigate how perception of a somatosensory stimulus differed according to whether it was delivered to a hand that was about to execute a reaching movement or the alternative, nonmoving, hand. The results of our study demonstrate that a somatosensory stimulus delivered to a hand that is being prepared for movement is perceived to have occurred later than when that same stimulus is delivered to a nonmoving hand. This result indicates that it takes longer for a tactile stimulus to be detected when it is delivered to a moving limb and may correspond to a change in perceptual threshold. Our behavioural results are paralleled by the results of our fMRI study that demonstrated that there were significantly reduced blood-oxygen-level-dependent (BOLD) responses within the parietal operculum and insula following somatosensory stimulation of the hand being prepared for movement, compared to when an identical stimulus was delivered to a nonmoving hand. These findings are consistent with the prediction of FM accounts of motor control that postulate that central sensory suppression of somatosensation accompanies self-generated limb movements, and with previous reports indicating that effects of sensory suppression are observed in higher order somatosensory regions.     

Exploring visuomotor priming following biological and non-biological stimuli

Observation of human actions influences the observer’s own motor system, termed visuomotor priming, and is believed to be caused by automatic activation of mirror neurons. Evidence suggests that priming effects are larger for biological (human) as opposed to non-biological (object) stimuli and enhanced when viewing stimuli in mirror compared to anatomical orientation. However, there is conflicting evidence concerning the extent of differences between biological and non-biological stimuli, which may be due to stimulus related confounds. Over three experiments, we compared how visuomotor priming for biological and non-biological stimuli was affected over views, over time and when attention to the moving stimulus was manipulated. The results indicated that the strength of priming for the two stimulus types was dependent on attentional location and load. This highlights that visuomotor priming is not an automatic process and provides a possible explanation for conflicting evidence regarding the differential effects of biological and non-biological stimuli.     

Matching of Movements Made Independently by the Two Arms in Normal Humans

Comparisons were made of voluntary movements of the right and left arms in normal human subjects. A series of movements of different amplitudes, made at the subject’s own speed, was performed with one limb. After a rest period, the same series was repeated with the contralateral limb. The relation between movement peak velocity and movement amplitude was linear and was the same for both arms. With repeated testing over periods up to two months, the slope of the peak velocity—amplitude relation decreased during the first week, thereafter remaining unchanged. In a second series of experiments, six normal subjects continuously wore a 1 lb (0.45 kg) weight strapped to their left (non-dominant) forearm for up to 1 week. This resulted in an increase in the slope of the peak-velocity/amplitude relation in this arm. A parallel change occurred in movements made independently by the right (non-loaded) arm. A similar matching of movement performance of the two limbs was seen following removal of the weight. The data is interpreted as providing support for the hypothesis that there is a single movement “command” which is applied to both limbs. The interaction of this command with the limbs which have similar second-order mechanical properties yields similar movements even when they are made independently.