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Human movement coordination implicates relative direction as the information for relative phase.

The current studies explore the informational basis of the coupling in human rhythmic movement coordination tasks. Movement stability in these tasks is an asymmetric U-shaped function of mean relative phase; 0 degrees is maximally stable, 90 degrees is maximally unstable and 180 degrees is intermediate. Bingham (2001, 2004a, 2004b) hypothesized that the information used to perform coordinated rhythmic movement is the relative direction of movement, the resolution of which is determined by relative speed. We used an experimental paradigm that entails using a circular movement to produce a linear motion of a dot on a screen, which must then be coordinated with a linearly moving computer controlled dot. This adds a component to the movement that is orthogonal to the display. Relative direction is not uniquely defined between orthogonal components of motion, but relative speed is; it was therefore predicted that the addition of the component would only introduce a symmetric noise component and not otherwise contribute to the U-shape structure of movement stability. Results for experiment 1 supported the hypothesis; movement that involved the additional component was overall less stable than movement that involved only the parallel component along which relative direction can be defined. Two additional studies ruled out alternative explanations for the pattern of data in experiment 1. Overall, the results strongly implicate relative direction as the information underlying performance in rhythmic movement coordination tasks.

Adult↗

The effect of visual transformation on bimanual circling movement.

In order to examine the effect of visual transformation on bimanual movements of various difficulty, fourteen participants performed bimanual circling tasks in three asymmetric movement modes--90 degrees (the left hand precedes the right hand by 1/4 cycle), 180 degrees (the delay between two hands is 1/2 cycle), and 270 degrees (the left hand precedes the right hand by 3/4 cycle)--under the normal vision condition and the visual transformation condition. In the visual transformation condition, movement of the right hand was transformed so that the required bimanual movement was always presented visually as a symmetric pattern. Additionally, the participants also performed a 0 degrees mode (in-phase symmetric) movement. Results revealed that the visual transformation increased the movement accuracy, with the variability of the right-left difference unchanged. Thus, proper visual transformation can improve the accuracy of a movement task. The 0 degrees mode was performed with higher stability and accuracy than any other movement modes of the visual transformation condition and normal vision conditions. In addition, the constant error associated with the 90 degrees and 270 degrees modes indicated that, in the normal vision condition, the executed movement was shifted to the 180 degrees mode, whereas in the visual transformation condition it stayed around the required mode and was slightly shifted to the 0 degrees mode. This result suggests that visual transformation can change the relationship between the intention to realize the required mode and the intrinsic neuromuscular dynamics. The effect size of visual transformation was larger in the 90 degrees and 270 degrees modes than in the 180 degrees mode. It is thus concluded that the effect of visual transformation depends upon the difficulty of the movement task.

Adult↗

Kinematic synergy adaptation to an unstable support surface and equilibrium maintenance during forward trunk movement.

The aim of this investigation was to study the adaptation to an unstable support surface of kinematic synergy responsible for equilibrium control during upper trunk movements. Eight adult subjects were asked to bend their upper trunk forward to an angle of 35 degrees and then to hold the final position for 3 s, first in a standard condition, with two feet on the ground and the second, on a rocking platform swinging in the sagittal plane. The movement characteristics (duration, amplitude, and mean angular velocity of the trunk), the time course of the antero-posterior center of mass (CM) shift during the movement, and the EMG pattern of the main muscles involved in the movement were studied under the two experimental conditions. Kinematic synergy was quantified by performing a principal component analysis on the hip, knee, and ankle angle changes occurring during the movement. The results indicate that (1) the CM shift from the very onset of the movement remains controlled during performance of the forward trunk movement when the equilibrium constraints were increased; (2) the principal component analysis of the hip, knee, and ankle angle changes occurring during the movement showed a transition from one principal component (PC(1)) in the standard condition to two components in the rocking platform condition; (3) the greatest contribution of PC(1) (weight coefficients) was located at the hip level in both the standard and rocking platform conditions, while the greatest contribution of PC(2) in the rocking platform condition was located at the ankle level; and (4) the EMG pattern underlying kinematic synergy is modified. It is concluded that a simple adaptation of kinematic synergy by changing the weight coefficients of each pair of joints participating in the movement is no longer sufficient when the equilibrium constraints increase and, rather, disturbs equilibrium. The CNS has to provide two parallel controls, one to perform the trunk movement and the other to preserve equilibrium.

Adaptation, Physiological↗

Eye movements cannot explain vibration-induced visual motion and motion aftereffect.

Eye movements are thought to account for a number of visual motion illusions involving stationary objects presented against a featureless background or apparent motion of the whole visual field. We tested two different versions of the eye movement account: (a) the retinal slip explanation and (b) the nystagmus-suppression explanation, in particular their ability to account for visual motion experienced during vibration of the neck muscles, and for the visual motion aftereffect following vibration. We vibrated the neck (ventral sternocleidomastoid muscles, bilaterally, or right dorsal muscles) and measured eye movements in conjunction with perceived illusory displacement of an LED presented in complete darkness (N=10). To test the retinal-slip explanation, we compared the direction of slow eye movements to the direction of illusory motion of the visual target. To test the suppression explanation, we estimated the direction of suppressed slow-phase eye movements and compared it to the direction of illusory motion. Two main findings show that neither actual nor suppressed eye movements cause the illusory motion and motion aftereffect. Firstly, eye movements do not reverse direction when the illusory motion reverses after vibration stops. Secondly, there are large individual differences with regards to the direction of eye movements in observers who all experience a similar visual illusion. We conclude that, rather than eye movements, a more global spatial constancy mechanism that takes into account head movement is responsible for the illusion. The results also argue against the notion of a single central signal that determines both perceptual experience and oculomotor behaviour.

Adult↗

Limits to human movement planning with delayed and unpredictable onset of needed information.

In motor tasks with explicit rewards and penalties, humans choose movement strategies that nearly maximize expected gain (Trommershäuser et al. in J Opt Soc Am A 20:1419-1433, 2003). Here, we examine whether performance is still close to optimal when information about payoffs is not available prior to movement onset. Subjects rapidly touched a target region while trying to avoid hitting an overlapping penalty region placed randomly to the left or right of the target. Subjects received rewards and incurred penalties for hitting the corresponding regions. Late responses (>700 ms) were heavily penalized. The penalty region was displayed 0, 200 or 400 ms after the reward region and the subject could not know where it would be until then. Reaction times to begin the movement after stimulus appearance were constant across conditions. Median reaction times were approximately 200 ms, i.e., the time the penalty was first displayed in the 200 ms delay condition. Performance was compared to that of an optimal movement planner that chooses mean end points to maximize expected gain despite movement variability. In the 0 and 200 ms delay conditions, subjects selected strategies that did not differ significantly from optimal, indicating that humans are able to plan their movements well despite delayed and unpredictable onset of information. Performance dropped below optimal in the 400 ms delay condition, with mean movement end points closer to the penalty region than predicted by the optimal strategy (in the high-penalty condition). We conclude that relevant information concerning the reward structure is required between 200 and 400 ms prior to the end of the movement, but can still be integrated into the movement plan after movement initiation.

Adult↗

Observation of a finger or an object movement primes imitative responses differentially.

Behavioural advantages for imitation of human movements over movements instructed by other visual stimuli are attributed to an 'action observation-execution matching' (AOEM) mechanism. Here, we demonstrate that priming/exogenous cueing with a videotaped finger movement stimulus (S1) produces specific congruency effects in reaction times (RTs) of imitative responses to a target movement (S2) at defined stimulus onset asynchronies (SOAs). When contrasted with a moving object at an SOA of 533 ms, only a human movement is capable of inducing an effect reminiscent of 'inhibition of return' (IOR), i.e. a significant advantage for imitation of a subsequent incongruent as compared to a congruent movement. When responses are primed by a finger movement at SOAs of 533 and 1,200 ms, inhibition of congruent or facilitation of incongruent responses, respectively, is stronger as compared to priming by a moving object. This pattern does not depend on whether S2 presents a finger movement or a moving object, thus effects cannot be attributed to visual similarity between S1 and S2. We propose that, whereas both priming by a finger movement and a moving object induces processes of spatial orienting, solely observation of a human movement activates AOEM. Thus, S1 immediately elicits an imitative response tendency. As an overt imitation of S1 is inadequate in the present setting, the response is inhibited which, in turn, modulates congruency effects.

Adult↗

Dissociation of motor preparation from memory and attentional processes using movement-related cortical potentials.

The EEG activity preceding self-paced voluntary movements (movement-related cortical potential, MRCP) is smaller if subjects make the same movement each time (regular task) compared with when different movements are made each time (random task). To test whether extra activity in the random task is due to increased motor preparation needed to switch between different movements, or to memory/attentional processes needed to select movements randomly, we compared regular and random movements with an additional alternating task. This alternating task required subjects to make different movements each time as in the random task, but since the task was very simple, the memory/attentional load was similar to that in the regular task. The MRCP was equally large over motor areas in both random and alternating tasks, suggesting that the extra activity over sensorimotor areas reflected processes involved in motor preparation rather than memory/attention. We speculate that, in the regular task, some part of the instructions for the previous movement remains intact, reducing the amount of preparation needed for the next repetition. Thus the MRCP is smaller than in the alternating and random tasks. Although the MRCPs in alternating and random tasks were similar over the motor areas, the random task had more activity than the alternating task in contralateral frontal areas. This part of the MRCP may therefore be related to memory/attentional processes required to randomize the sequence of movements. We conclude that the MRCP contains dissociable components related to motor preparation and memory/attention.

Adult↗

Movement discrimination by single cells in the human pallidum characterised by hidden Markov models.

In patients undergoing pallidotomy for Parkinson's disease, we recorded extracellularly from single neurons in the two internal segments (GPii, GPie) and the external segment (GPe) of the globus pallidus (GP) in response to active (cued) movements of the contralateral wrist, elbow or ankle. The patterns of cell activity occurring both before and after movement onset were analysed using hidden Markov models (HMMs) and clustered by movement type using the generative topographical mapping algorithm. Cluster separation was quantified in order to measure a cell's ability to discriminate between movements. Statistical analysis of variance indicated a significant regional gradient (GPii > GPie > GPe) of movement discrimination, while cells in all regions differentiated better between movements of different joints (wrist, elbow or ankle) than between flexion and extension of the same joint. We found that GP cells generally showed distinguishable firing patterns corresponding to more than one type of movement per cell, in support of the hypothesis that cells in these regions of the basal ganglia are not involved in preparation or execution of a single type of movement but participate in many different movements, analogous to the hidden units of a neural network. Our results also indicate that cell activity both preceding a movement and during its execution may be modelled by HMMs with only a small number of states.

Action Potentials↗

Microstimulation of movements from cerebellar-receiving, but not pallidal-receiving areas of the macaque thalamus under ketamine anaesthesia.

The motor thalamic areas receiving input from the globus pallidus (VA) and the cerebellar nuclei (VL) appear to have different roles in the generation and guidance of movements. In order to further test these differences, we used electrical stimulation to map the ventro-anterior and ventro-lateral nuclei of the thalamus in three ketamine anaesthetised monkeys. Movements were readily evoked from VL at currents of down to 10 microA. The movements were typically multijoint, and stimulation could evoke arm and trunk or arm and facial movement at the same current threshold. Evoked arm movements often involved multiple joints, with or without finger movements. Facial movements included the lips, tongue, jaw, eyebrows and, occasionally, the eyes. The thalamic map was topographic, but complex with at least two separate regions related to arm movement. Very few sites within the VA could stimulate movement, even at high currents. We therefore suggest that the cerebellar projections to motor regions of the cortex, which pass through the VL thalamic nuclei, have a different relationship and are closer to movement execution than the projections from basal ganglia via the ventro-anterior nucleus.

Anesthetics, Dissociative↗

PET study of visually and non-visually guided finger movements in patients with severe pan-sensory neuropathies and healthy controls.

Although patients with sensory neuropathies and normal muscle power are rare, they have been extensively studied because they are a model for dissociating the sensory and motor components of movement. We have examined these patients to determine the cerebral functional anatomy of movement in the absence of proprioceptive input. In addition, the disabling symptoms of these patients can be substantially improved by visually monitoring their movements. We hypothesized that, during visually guided movements, these patients would show overactivity of regions specialized for visuomotor control with the possible additional involvement of areas that normally process somatosensory information. We used positron emission tomography (PET) and the tracer H(2)(15)O to determine the functional anatomy of visually and non-visually guided finger movements in three patients with long-standing pan-sensory neuropathies and normal muscle power and six healthy controls. Five conditions were performed with the right hand: a sequential finger movement task under visual guidance, the same motor task without observation of the hand, monitoring a video of the same sequential finger movement, a passive visual task observing a reversing checkerboard, and an unconstrained rest condition. Data were analyzed using conventional subtraction techniques with a statistical threshold of z>2.33 with corrections for multiple comparisons. When compared with the control group, activation was not deficient in any brain areas of the patient cohort in any of the contrasts tested. In particular, in the non-visually guided movement task, in which meaningful visual and proprioceptive input was absent, the patient group activated primary motor, premotor, and cerebellar regions. This suggests that these areas are involved in motor processing independent of sensory input. In all conditions involving visual observation of hand movements, there was highly significant overactivity of the left parietal operculum (SII) and right parieto-occipital cortex (PO) in the patient group. Recent non-human primate studies have suggested that the PO region contains a visual representation of hand movements. Overactivity of this area and the activation of SII by visual input appear to indicate that compensatory overactivity of visual areas and cross-modal plasticity of somatosensory areas occur in deafferented patients. These processes may underlie their ability to compensate for their proprioceptive deficits.

Aged↗

On the role of static and dynamic visual afferent information in goal-directed aiming movements.

Movement planning has been shown to be optimized when the participant is permitted to see his or her hand resting on the starting base prior to movement initiation. However, this proposition is opposed by contradictory results. In the present study, we wanted to determine whether these conflicting results were caused by procedural differences. The results showed that seeing one's hand on the starting base did not result in more accurate aiming movement than when this information was not available. However, lower aiming errors were found when one was asked to foveate the starting base and then the target prior to movement initiation, but only when no dynamic visual information was available during movement. When an aiming movement was performed while one's hand was visible in visual periphery, foveating the starting base or not prior to movement initiation did not modify aiming accuracy. These results suggest that gazing at the starting base and then at the target provides an eye-based representation of the movement to be performed that can be used by the CNS to plan a manual aiming movement. Information for better planning of the direction--but not the extent--dimension of an upcoming movement can also be derived from dynamic visual information available in peripheral vision.

Adult↗

Assessment of movement time in patients with essential tremor.

BACKGROUND: We have observed mild bradykinesia in essential tremor (ET) patients, which do not satisfy the criteria of Parkinson's disease (PD). OBJECTIVE: To compare the mean movement time for repetitive movements around distal and proximal joints in ET patients with normal controls by using a simple test paradigm. PATIENTS AND METHODS: Seventeen patients with ET and 14 control subjects were instructed to tap with the index finger sequentially on push-button microswitches. Movement times around metacarpophalangeal, wrist, elbow, and shoulder joints of the right side were tested. The data collected were stored on a computer and the time elapsed between sequential taps on two keys (ms) and number of taps on the left key for 15 seconds were evaluated offline. RESULTS: Movement times of the patients with ET were not found to be significantly different from those of the controls at all joints tested despite slight prolongation for movements around the shoulder joint. CONCLUSION: The simple test paradigm we have used showed that there is no difference in the movement time for repetitive movements around four joints of the upper extremity between patients with ET and normal control subjects. The slightly prolonged movement time around the shoulder joint noted in patients with ET may be ascribed to tremor, not bradykinesia. Tremor may cause these patients to pay more attention to the performance of goal-directed finger movements and consequently prolong movement time slightly or it may simply delay the time elapsed to reach the goal in the absence of overt intention tremor.

Aged↗

A numerical simulation of muscle spindle ensemble encoding during planar movement of the human arm.

We address the issue of what proprioceptive information, regarding movement of the human arm, may be provided to the central nervous system by proprioceptors located within muscles of this limb. To accomplish this we developed a numerical simulation which could provide estimates of the length regimes experienced by a set of model receptors located within some of the principal muscles of the human arm during planar movement of this limb. These receptors were assumed to have characteristics analogous to those associated with a simple model of muscle spindle signalling of movement. To this end each spindle had proprioceptive 'channels' associated with it. These corresponded to primary and secondary spindle afferent fibers which could provide independent afferent output regarding the parent muscle the spindle monitored. The angles of the shoulder and elbow joints attained by subjects performing a task requiring movement of the right arm in a horizontal plane to a static visual target were recorded. For this angular data the lengths and rates of change of lengths experienced by muscle fascicles, and hence the model spindles, during movement were calculated by means of the numerical simulation. The discharge rates of the simulated spindles during the movement were calculated to derive a measure of the depth of modulation, induced by the movement, for each spindle. These values were then summed for all spindles to provide a first-order approximation of spindle ensemble coding of the movement. Significant correlations (P < 0.0001, Spearman's rank order) were found between the resulting ensemble encodings and, in order of significance, the angular velocity of the shoulder joint (rS = 0.945), the tangential velocity of the hand (rS = 0.942), and the angular velocity of the elbow joint (rS = 0.917). Correlations between the angular positions of the shoulder (rS = -0.623) and elbow (rS = 0.628) were lower. These findings indicate that the ensemble profiles of the simulated muscle spindles, encode information regarding kinematic parameters of movements related to both intrinsic and extrinsic coordinate systems. This suggests that motor structures capable of deriving such an ensemble encoding would be in a position to perform the sensory-motor transformations between intrinsic and extrinsic frames of reference necessary for controlling movements planned in extrinsic coordinates.

Arm↗

Does the brain use sliding variables for the control of movements?

Delays in the transmission of sensory and motor information prevent errors from being instantaneously available to the central nervous system (CNS) and can reduce the stability of a closed-loop control strategy. On the other hand, the use of a pure feedforward control (inverse dynamics) requires a perfect knowledge of the dynamic behavior of the body and of manipulated objects. Sensory feedback is essential both to accommodate unexpected errors and events and to compensate for uncertainties about the dynamics of the body. Experimental observations concerning the control of posture, gaze and limbs have shown that the CNS certainly uses a combination of closed-loop and open-loop control. Feedforward components of movement, such as eye saccades, occur intermittently and present a stereotyped kinematic profile. In visuo-manual tracking tasks, hand movements exhibit velocity peaks that occur intermittently. When a delay or a slow dynamics are inserted in the visuo-manual control loop, intermittent step-and-hold movements appear clearly in the hand trajectory. In this study, we investigated strategies used by human subjects involved in the control of a particular dynamic system. We found strong evidence for substantial nonlinearities in the commands produced. The presence of step-and-hold movements seemed to be the major source of nonlinearities in the control loop. Furthermore, the stereotyped ballistic-like kinematics of these rapid and corrective movements suggests that they were produced in an open-loop way by the CNS. We analyzed the generation of ballistic movements in the light of sliding control theory assuming that they occurred when a sliding variable exceeded a constant threshold. In this framework, a sliding variable is defined as a composite variable (a combination of the instantaneous tracking error and its temporal derivatives) that fulfills a specific stability criterion. Based on this hypothesis and on the assumption of a constant reaction time, the tracking error and its derivatives should be correlated at a particular time lag before movement onset. A peak of correlation was found for a physiologically plausible reaction time, corresponding to a stable composite variable. The direction and amplitude of the ongoing stereotyped movements seemed also be adjusted in order to minimize this variable. These findings suggest that, during visually guided movements, human subjects attempt to minimize such a composite variable and not the instantaneous error. This minimization seems to be obtained by the execution of stereotyped corrective movements.

Adult↗

Impedance characteristics of a neuromusculoskeletal model of the human arm II. Movement control.

The modulation of neuromusculoskeletal impedance during movements is analysed using a motor control model of the human arm. The motor control system combines feedback and feedforward control and both control modes are determined in one optimization process. In the model, the stiffness varies at the double movement frequency for 2-Hz oscillatory elbow movements and has high values at the movement reversals. During goal-directed two-degrees-of-freedom arm movements, the stiffness is decreased during the movement and may be increased in the initial and final phases, depending on the movement velocity. The stiffness has a considerable curl during the movement, as was also observed in experimental data. The dynamic stiffness patterns of the model can be explained basically by the alpha-gamma coactivation scheme where feedback gains covary with motor control signals. In addition to the modulation of the gain factors, it is argued that the variation of the intrinsic stiffness has a considerable effect on movement control, especially during fast movements.

Arm↗

Breech presentation: is there a difference in eye movement patterns compared with cephalic presentation in the human fetus at term?

OBJECTIVE: Our purpose was to characterize eye movement patterns in the human fetus at term persisting in breech presentation. STUDY DESIGN: Studied were 11 fetuses in breech presentation and 12 in cephalic presentation at 36 to 41 weeks' gestation, of which the presentation remained unchanged from 32 weeks' gestation until delivery. With real-time ultrasonography we analyzed three measurements by means of the Mann-Whitney rank-sum test: (1) duration of eye-movement and no-eye-movement periods, (2) frequency of eye movements, and (3) proportion of horizontal, vertical, and oblique eye movements. RESULTS: There were significant differences in the eye movement directions between breech and cephalic fetuses: horizontal (median 70.3% vs 80.2%), vertical (17.4% vs 11.0%), and oblique (13.1% vs 9.1%), respectively. No differences were noted in the duration of eye-movement and no-eye-movement periods or in the frequency of eye movements. CONCLUSION: Eye movement pattern, in direction, of term fetuses persisting in breech presentation run on a different in-utero developmental course of neural control from those in cephalic fetuses.

Breech Presentation↗

A computer model of jaw and hyoid movement in the cat.

A digital model was used to study the varying movements of the feline jaw and hyoid in the sagittal plane. The effect of mandibular movement on the path of movement of the hyoid relative to the palate was investigated for model paths of movement of the hyoid, i.e. paths of movement that were always elliptical relative to the mandible. Major effects upon the path of movement of the hyoid relative to the palate were produced by two factors: (1) the presence of a fast open/fast close complex within the profile of jaw movement; (2) the time relationship of the backwards movement of the hyoid relative to that of the fast open/fast close complex. The average antero-posterior position of the hyoid in the mandibular plane and the angle of the major axis of the ellipse to that plane were less important factors. Within limits (relative to the mandible), the overall amplitude of the hyoid movement, its vertical position and the direction in which the ellipse was travelled were all unimportant factors in determining the general nature of hyoid movement relative to the palate.

Animals↗

Human jaw movement in mastication and speech.

The study of jaw movement in humans is a primary source of information about the relationship between voluntary movement and more primitive motor functions. This study focused on the geometric form of the velocity function, as measured by linear voltage displacement transducer. Movement amplitudes, maximum velocities and durations were greater in mastication than in speech. Nevertheless, there were detailed similarities in the shape of the normalized velocity functions. In jaw-closing movements, the normalized functions were similar in form over differences in rate, movement amplitude (speech movements) and the compliance of the bolus (mastication). In opening movements, the functions for mastication and speech were again similar over differences in amplitude and compliance. However, they differed in shape for fast and slow movements. Normalized acceleration and deceleration durations were approximately equal in rapid movements, whereas, for slower movements, deceleration took substantially longer than acceleration.

Biomechanical Phenomena↗