Motor synergies for dampening hand vibration during human walking

This study investigated the motion required to carry a cup filled with water without spilling it, which is a common human dexterous task. This task requires the individual to dampen hand vibration while walking. We hypothesize that a reduction in hand jerk and a constant cup angle are required to achieve this task. We measured movements while human subjects carried a cup with water (WW task) and with stones (WS task) using a three-dimensional position measurement system and then analyzed joint coordination. We empirically confirmed that the value of hand jerk and the variance in cup angle in the WW task were smaller than those in the WS task. We used uncontrolled manifold (UCM) analysis to quantify joint coordination corresponding to the motor synergy required to reduce the hand jerk and variance of the cup angle. UCM components, which did not affect the hand jerk and cup angle, were larger than orthogonal components, which directly affected the hand jerk and cup angle in the WW task. These results suggest that there is a coordinated control mechanism that reduces hand jerk and maintains a constant cup angle when carrying a cup filled with water without spilling it. In addition, we suggest that humans adopt a flexible and coordinated control strategy of allowing variance independent of the variables that should be controlled to achieve this dexterous task.     

Joint angle variability and co-variation in a reaching with a rod task

The problem at the heart of motor control is how the myriad units of the neuromotor system are coordinated to perform goal-directed movements. Although for long these numerous degrees of freedom (DOFs) were considered redundant, recent views emphasize more that the DOFs should be considered abundant, allowing flexible performance. We studied how variability in arm joints was employed to stabilize the displaced end-effector in tool use to examine how the neuromotor system flexibly exploits DOFs in the upper extremity. Participants made pointing movements with the index finger and with the index finger extended by rods of 10, 20, and 30 cm. Using the uncontrolled manifold (UCM) method, the total joint angle variance was decomposed into two parts, the joint angle variance that did not affect the position of the end-effector (V _UCM) and the variance that results in a deviation of the position of the end-effector from its mean (V _ORT). Analyses showed that some angles depended on length of the rod in use. For all rod lengths, V _UCM was larger than V _ORT, and this did not differ over rod lengths, demonstrating that the arm was organized into a synergy. Finally, the variation in the joint angles in the arm as well as the degree of co-variation between these angles did not differ for the rod’s tip and the hand. We concluded that synergies are formed in the arm during reaching with an extended end-effector and those synergies stabilize different parts of the arm+rod system equally.     

Motor equivalent control of the center of mass in response to support surface perturbations

To claim that the center of mass (CM) of the body is a controlled variable of the postural system is difficult to verify experimentally. In this report, a new variant of the method of the uncontrolled manifold (UCM) hypothesis was used to evaluate CM control in response to an abrupt surface perturbation during stance. Subjects stood upright on a support surface that was displaced in the posterior direction. Support surface translations between 0.03 and 0.12 m, each lasting for 275 ms, were presented randomly. The UCM corresponding to all possible combinations of joints that are equivalent with respect to producing the average pre-perturbation anterior–posterior position of the center of mass (CM_AP) were linearly estimated for each trial. At each point in time thereafter, the difference between the current joint configuration and the average pre-perturbation joint configuration was computed. This joint difference vector was then projected onto the pre-perturbation UCM as a measure of motor equivalence, and onto its complementary subspace, which represents joint combinations that lead to a different CM_AP position. A similar analysis was performed related to control of the trunk’s spatial orientation. The extent to which the joint velocity vector acted to stabilize the CM_AP position was also examined. Excursions of the hip and ankle joints both increased linearly with perturbation magnitude. The configuration of joints at each instance during the perturbation differed from the mean configuration prior to the perturbation, as evidenced by the joint difference vector. Most of this joint difference vector was consistent, however, with the average pre-perturbation CM_AP position rather than leading to a different CM_AP position. This was not the case, however, when performing this analysis with respect to the UCM corresponding to the control of the pre-perturbation trunk orientation. The projection of the instantaneous joint velocity vector also was found to lie primarily in the UCM corresponding to the pre-perturbation CM_AP position, indicating that joint motion was damped in directions leading to a change away from the pre-perturbation CM_AP position. These results provide quantitative support for the argument that the CM position is a planned variable of the postural system and that its control is achieved through selective, motor equivalent changes in the joint configuration in response to support surface perturbations. The results suggest that the nervous system accomplishes postural control by a control strategy that considers all DOFs. This strategy presumably resists combinations of DOFs that affect the stability of important task-relevant variables (CM_AP position) while, to a large extent, freeing from control combinations of those DOFs that have no effect on the task-relevant variables (Schöner in Ecol Psychol 8:291–314, 1995).     

How visual information links to multijoint coordination during quiet standing

The link between visual information and postural control was investigated based on a multi-degree-of-freedom model using the framework of the uncontrolled manifold (UCM) hypothesis. The hypothesis was that because visual information specifies the position of the body in space, it would couple preferentially into those combinations of degrees of freedom (DOFs) that move the body in space and not into combinations of DOFs that do not move the body in space. Subjects stood quietly in a virtual reality cave for 4-min trials with or without a 0.2, 2.0 Hz, or combined 0.2 and 2.0 Hz visual field perturbation that was below perceptual threshold. Motion analysis was used to compute six sagittal plane joint angles. Variance across time of the angular motion was partitioned into (1) variance associated with motion of the body and (2) variance reflecting the use of flexible joint combinations that keep the anterior–posterior positions of the head (HD_POS) and center of mass (CM_POS) invariant. UCM analysis was performed in the frequency domain in order to link the sensory perturbation to each variance component at different frequencies. As predicted, variance related to motion of the body was selectively increased at the 0.2-Hz drive frequency but not at other frequencies of sway for both CM_POS and HD_POS. The dominant effect with the 2.0-Hz visual drive also was limited largely to variance related to motion of the body.     

The dynamics of postural sway cannot be captured using a one-segment inverted pendulum model: a PCA on segment rotations during unperturbed stance

Research on unperturbed stance is largely based on a one-segment inverted pendulum model. Recently, an increasing number of studies report a contribution of other major joints to postural control. Therefore this study evaluates whether the conclusions originating from the research based on a one-segment model adequately capture postural sway during unperturbed stance. High-pass filtered kinematic data (cutoff frequency 1/30 Hz) obtained over 3 min of unperturbed stance were analyzed in different ways. Variance of joint angles was analyzed. Principal-component analysis (PCA) was performed on the variance of lower leg, upper leg, and head-arms-trunk (HAT) angles, as well as on lower leg and COM angle (the orientation of the line from ankle joint to center of mass). It was found that the variance in knee and hip joint angles did not differ from the variance found in the ankle angle. The first PCA component indicated that, generally, the upper leg and HAT segments move in the same direction as the lower leg with a somewhat larger amplitude. The first PCA component relating ankle angle variance and COM angle variance indicated that the ankle joint angle displacement gives a good estimate of the COM angle displacement. The second PCA component on the segment angles partly explains the apparent discrepancy between these findings because this component points to a countermovement of the HAT relative to the ankle joint angle. It is concluded that postural control during unperturbed stance should be analyzed in terms of a multiple inverted pendulum model.     

Joint coordination during quiet stance: effects of vision

Stabilization of the center of mass (CM) is an important goal of the postural control system. Coordination of several joints along the human pendulum is required to achieve this goal. We studied the coordination among body segments with respect to horizontal CM stabilization during a quiet stance task and the effects of vision on CM stability. Subjects were asked to stand quietly on a narrow wooden block supporting only the mid-foot, with either open (EO) or closed (EC) eyes on separate trials. Instant equilibrium points (IEPs) in the center of pressure (CP) trajectory were determined when the horizontal component of the ground reaction force was zero and the CP data were decomposed into their rambling and trembling components. The joint angle, CM and CP data were divided into short cycles (time-normalized to 100 data points) or longer segments (time-normalized to 1000 data points) of equal length beginning and ending in an IEP. Motor abundance with respect to patterns of joint coordination was evaluated using the uncontrolled manifold (UCM) approach. Here, a UCM is a subspace spanning all joint combinations resulting in a given CM position. All combinations of joint angles that lie within this subspace are equivalent with respect to that CM position while joint angle combinations lying in a subspace orthogonal to the UCM lead to deviation from that CM position. UCM analysis was performed on data organized either across time within longer segments or at each point in time across multiple segments or across multiple cycles. Regardless of method of analysis, most of the variance in joint space was constrained to be within the UCM, preserving the mean CM position in both the EO and EC conditions. Joint configuration variance was significantly higher in the EC than in the EO condition although this increase occurred primarily within the UCM rather than in the orthogonal subspace that would have led to variation of the CM position. These results demonstrate the ability of the control system to selectively channel motor variability into directions in joint space that stabilize the CM position. This effect was enhanced when the task was made more challenging in the absence of vision. There was also a significant relationship between joint variance that led to a change in the CM position and, in particular, the rambling component of the CP path, lending some support to the idea that the CNS prescribes a certain stable trajectory of the CP during quiet stance that leads to a small controlled movement of the CM.     

Reference frames for spinal proprioception: limb endpoint based or joint-level based?

Many sensorimotor neurons in the CNS encode global parameters of limb movement and posture rather than specific muscle or joint parameters. Our investigations of spinocerebellar activity have demonstrated that these second-order spinal neurons also may encode proprioceptive information in a limb-based rather than joint-based reference frame. However, our finding that each foot position was determined by a unique combination of joint angles in the passive limb made it difficult to distinguish unequivocally between a limb-based and a joint-based representation. In this study, we decoupled foot position from limb geometry by applying mechanical constraints to individual hindlimb joints in anesthetized cats. We quantified the effect of the joint constraints on limb geometry by analyzing joint-angle covariance in the free and constrained conditions. One type of constraint, a rigid constraint of the knee angle, both changed the covariance pattern and significantly reduced the strength of joint-angle covariance. The other type, an elastic constraint of the ankle angle, changed only the covariance pattern and not its overall strength. We studied the effect of these constraints on the activity in 70 dorsal spinocerebellar tract (DSCT) neurons using a multivariate regression model, with limb axis length and orientation as predictors of neuronal activity. This model also included an experimental condition indicator variable that allowed significant intercept or slope changes in the relationships between foot position parameters and neuronal activity to be determined across conditions. The result of this analysis was that the spatial tuning of 37/70 neurons (53%) was unaffected by the constraints, suggesting that they were somehow able to signal foot position independently from the specific joint angles. We also investigated the extent to which cell activity represented individual joint angles by means of a regression model based on a linear combination of joint angles. A backward elimination of the insignificant predictors determined the set of independent joint angles that best described the neuronal activity for each experimental condition. Finally, by comparing the results of these two approaches, we could determine whether a DSCT neuron represented foot position, specific joint angles, or none of these variables consistently. We found that 10/70 neurons (14%) represented one or more specific joint-angles. The activity of another 27 neurons (39%) was significantly affected by limb geometry changes, but 33 neurons (47%) consistently elaborated a foot position representation in the coordinates of the limb axis.     

Control and estimation of posture during quiet stance depends on multijoint coordination

This study tested the hypotheses that all major joints along the longitudinal axis of the body are equally active during quiet standing and that their motions are coordinated to stabilize the spatial positions of the center of mass (CM) and head. Analyses of the effect of joint configuration variance on the stability of the CM and head positions were performed using the uncontrolled manifold (UCM) approach. Subjects stood quietly with arms folded across their chests for three 5-min trials each with and without vision. The UCM analysis revealed that the six joints examined were coordinated such that their combined variance had minimal effect on the CM and head positions. Removing vision led to a structuring of the resulting increased joint variance such that little of the increase affected stability of the CM and head positions. The results reveal a control strategy involving coordinated variations of most major joints to stabilize variables important to postural control during quiet stance.     

The effect of movement direction on joint torque covariation

It has been proposed that unconstrained upper limb movements are coordinated via a kinetic constraint that produces dynamic muscle torques at each moving joint that are a linear function of a single torque command. This constraint has been termed linear synergy (Gottlieb et al. J Neurophysiol 75:1760–1764, 1996). The current study tested two hypotheses: (1) that the extent of covariation between dynamic muscle torques at the shoulder and elbow varied with the direction of movement and (2) that the extent to which muscle torques deviated from linear synergy would be reproduced by a simulation of pointing movements in which the path of the hand was constrained to be straight. Dynamic muscle torques were calculated from sagittal plane pointing movements performed by 12 participants to targets in eight different directions. The results of principal component analyses performed on the muscle torque data demonstrated direction-dependent variation in the extent to which dynamic muscle torques covaried at the shoulder and elbow. Linear synergy was deviated from substantially in movement directions for which the magnitude of muscle torque was low at one joint. A simulation of movements with straight hand paths was able to accurately estimate the amount of covariation between muscle torques at the two joints in many directions. These results support the idea that a kinematic constraint is imposed by the central nervous system during unconstrained pointing movements. Linear synergy may also be applied as a coordinating constraint in circumstances where its application allows the path of the moving endpoint to remain close to a straight line.     

Kinematic and kinetic constraints on arm, trunk, and leg segments in target-reaching movements

We studied target reaching tasks involving not only the arms but also the trunk and legs, which necessitated some trunk flexion. Such tasks can be successfully completed using an infinite number of combinations of segment motions due to the inherent kinematic redundancy with the excessive degrees of freedom (DOFs). Sagittal plane motions of six segments (shank, thigh, pelvis, trunk, humerus, and forearm) and dynamic torques of six joints (ankle, knee, hip, lumbar, shoulder, and elbow) were analyzed separately by principal component (PC) analyses to determine if there was a commonality among the shapes of the respective waveforms. Additionally, PC analyses were used to probe for constraining relationships among the 1) relative magnitudes of segment excursions and 2) the peak-to-peak dynamic joint torques. In summary, at the kinematic level, the tasks are simplified by the use of a single common waveform for all segment excursions with 89.9% variance accounted for (VAF), but with less fixed relationships among the relative scaling of the magnitude of segment excursions (62.2% VAF). However, at the kinetic level, the time course of the dynamic joint torques are not well captured by a single waveform (72.7% VAF), but the tasks are simplified by relatively fixed relationships among the scaling of dynamic joint torque magnitudes across task conditions (94.7% VAF). Taken together, these results indicate that, while the effective DOFs in a multi-joint task are reduced differently at the kinematic and kinetic levels, they both contribute to simplifying the neural control of these tasks.     

Learning a throwing task is associated with differential changes in the use of motor abundance

This study sought to characterize changes in the synergy of joint motions related to learning a Frisbee throwing task and, in particular, how the use of abundant solutions to joint coordination changed during the course of learning for successful performance. The latter information was helpful in determining the relative importance of different performance-related variables (PVs) to performance improvement. Following a pre-test, the main experiment consisted of six subjects practicing a Frisbee throw to a laterally-placed target for five days, 150 throws per day, followed by a post-test. A subgroup of three subjects continued to practice for an extended period of extensive practice amounting to 1800–2700 additional throws each, followed by a second post-test. Motor abundance was addressed through the uncontrolled manifold approach (UCM), which was used to partition the variance of joint configurations into two components with respect to relevant PVs, one component leading to a consistent value of the PV across repetitions, and a reflection of motor abundance, and a second component resulting in unstable values of the relevant PV. The method was used to test hypotheses about the relative importance of controlling the PVs that have an impact on successful task performance: movement extent, movement direction, hand path velocity, and the hands orientation to the target. In addition, the amount of self-motion, or apparently extraneous joint motion having no effect on the hands motion, compared to joint motion that does affect the hands motion, was determined. After a week of practice, all subjects showed improvement in terms of targeting accuracy. Hand movement variability also decreased with practice and this was associated with a decrease in overall joint configuration variance. This trend continued to a greater extent in the three subjects who participated in extended practice. Although the component of joint configuration variance that was consistent with a stable value of all PVs was typically substantially higher than variance leading to unstable values of those PVs, both components decreased with practice. However, the decrease in joint configuration variance reflecting motor abundance was less than the other variance component only in relation to control of movement direction and the hands orientation to the target. These results indicate that improvement of throwing performance in this experiment was more related to improved stabilization of movement direction and to the hands orientation to the target than to movement extent and hand velocity. Nonetheless, the relative values of the two joint variance components were such that the instantaneous value of both hand path velocity and movement extent were stabilized throughout the experiment and showed a consistent compensatory relationship at the time of Frisbee release, despite not changing with practice. Finally, the amount of self-motion increased significantly with practice, possibly reflecting better compensation for perturbations due to the limbs dynamics. The results are consistent with other studies, suggesting the need to reevaluate Bernsteins hypothesis of freeing and freezing DOFs with learning.     

男性青年踝关节外侧副韧带损伤后行走步态分析

目的 获取青年男性踝关节外侧韧带损伤者行走时踝关节的运动学和动力学参数,探讨损伤者行走步态的生物力学特性。方法 采用Qualisys MCU500三维运动影像捕捉系统与Kistler三维测力台,对踝关节外侧副韧带损伤者与健康者各15名的行走步态进行三维同步测试。结果 踝关节外侧副韧带损伤者竖直方向的地面反力变化平缓;在支撑时相中前期,损伤者前后方向的力要比健康者大;在步态周期的60%之前,损伤者左右方向的力明显大于健康者。损伤者关节跖屈力矩变化与健康者相似,患侧外翻力矩、外旋力矩最大,损伤者健侧输出功率最大。结论 损伤者踝关节的稳定性下降,行走时步态异常,为减少患侧负荷,健侧出现代偿效应,足着地瞬间加速过度到垂直支撑时相。本研究为预防踝关节以及损伤后的临床治疗以及康复训练等提供一定的理论参考。     

Multijoint error compensation mediates unstable object control

A key feature of skilled object control is the ability to correct performance errors. This process is not straightforward for unstable objects (e.g., inverted pendulum or "stick" balancing) because the mechanics of the object are sensitive to small control errors, which can lead to rapid performance changes. In this study, we have characterized joint recruitment and coordination processes in an unstable object control task. Our objective was to determine whether skill acquisition involves changes in the recruitment of individual joints or distributed error compensation. To address this problem, we monitored stick-balancing performance across four experimental sessions. We confirmed that subjects learned the task by showing an increase in the stability and length of balancing trials across training sessions. We demonstrated that motor learning led to the development of a multijoint error compensation strategy such that after training, subjects preferentially constrained joint angle variance that jeopardized task performance. The selective constraint of destabilizing joint angle variance was an important metric of motor learning. Finally, we performed a combined uncontrolled manifold-permutation analysis to ensure the variance structure was not confounded by differences in the variance of individual joint angles. We showed that reliance on multijoint error compensation increased, whereas individual joint variation (primarily at the wrist joint) decreased systematically with training. We propose a learning mechanism that is based on the accurate estimation of sensory states.     

Reciprocal angular acceleration of the ankle and hip joints during quiet standing in humans

Human quiet standing is often modeled as a single inverted pendulum rotating around the ankle joint, under the assumption that movement around the hip joint is quite small. However, several recent studies have shown that movement around the hip joint can play a significant role in the efficient maintenance of the center of body mass (COM) above the support area. The aim of this study was to investigate how coordination between the hip and ankle joints is controlled during human quiet standing. Subjects stood quietly for 30 s with their eyes either opened (EO) or closed (EC), and we measured subtle angular displacements around the ankle (thetaa) and hip (thetah) joints using three highly sensitive CCD laser displacement sensors. Reliable data were obtained for both angular displacement and angular velocity (the first derivative of the angular displacement). Further, measurement error was not predominant, even among the angular acceleration data, which were obtained by taking the second derivative of the angular displacement. The angular displacement, velocity, and acceleration of the hip were found to be significantly greater (P<0.001) than those of the ankle, confirming that hip-joint motion cannot be ignored, even during quiet standing. We also found that a consistent reciprocal relationship exists between the angular accelerations of the hip and ankle joints, namely positive or negative angular acceleration of ankle joint is compensated for by oppositely directed angular acceleration of the hip joint. Principal component analysis revealed that this relationship can be expressed as: thetah=gammathetaa with gamma=-3.15+/-1.24 and gamma=-3.12+/-1.46 (mean +/-SD) for EO and EC, respectively, where theta is the angular acceleration. There was no significant difference in the values of y for EO and EC, and these values were in agreement with the theoretical value calculated assuming the acceleration of COM was zero. On the other hand, such a consistent relationship was never observed for angular displacement itself. These results suggest that the angular motions around the hip and ankle joints are not to keep the COM at a constant position, but rather to minimize acceleration of the COM.     

What do synergies do? Effects of secondary constraints on multidigit synergies in accurate force-production tasks

We used the framework of the uncontrolled manifold (UCM) hypothesis to explore changes in the structure of variability in multifinger force-production tasks when a secondary task was introduced. Healthy young subjects produced several levels of the total force by pressing with the four fingers of the hand on force sensors. The frame with the sensors rested on the table (Stable condition) or on a narrow supporting beam (Unstable conditions) that could be placed between different finger pairs. Most variance in the finger mode space was compatible with a fixed value of the total force across all conditions, whereas the patterns of sharing of the total force among the fingers were condition dependent. Moment of force was stabilized only in the Unstable conditions. The finger mode data were projected onto the UCM computed for the total force and subjected to principal component (PC) analysis. Two PCs accounted for >90% of the variance. The directions of the PC vectors varied across subjects in the Stable condition, whereas two "default" PCs were observed under the Unstable conditions. These observations show that different persons coordinate their fingers differently in force-production tasks. They converge on similar solutions when an additional constraint is introduced. The use of variable solutions allows avoiding a loss in accuracy of performance when the same elements get involved in another task. Our results suggest a mechanism underlying the principle of superposition suggested in a variety of human and robotic studies.     

Effects of varying task constraints on solutions to joint coordination in a sit-to-stand task

The question of how multijoint movement is controlled can be studied by discovering how the variance of joint trajectories is structured in relation to important task-related variables. In a previous study of the sit-to-stand task, for instance, variations of body segment postures that leave the position of the body's center of mass (CM) unchanged were significantly greater than variations of body segment posture that varied the CM position. The present experiments tested the hypothesis that such structuring of joint configuration variability is accentuated when the mechanical or perceptual task demands are made more challenging. Six subjects performed the sit-to-stand task without vision (eyes closed), either on a normal or on a narrow support surface. An additional constraint on the postural task was introduced in a third condition by requiring subjects to maintain light touch (less than 1 N) with the fingertips while coming to a standing position on the narrow base of support. The joint configurations observed at each point in normalized time were analyzed with respect to trial-to-trial variability. The task variables CM and head position were used to define goal-equivalent sets of joint configurations ("uncontrolled manifolds," UCMs) within which variation of joint configuration leaves the task variables unchanged. The variability of joint configurations across trials was decomposed into components that did not affect (within the UCM) and that did affect (orthogonal to the UCM) the values of these task variables. Our results replicate the earlier finding of much larger variability in directions of joint space that leave the CM unchanged compared with directions that affect CM position. This effect was even more pronounced here than in the previous experiment, probably because of the more difficult perceptual conditions in the current study (eyes closed). When the mechanical difficulty of the task was increased, the difference between the two types of joint variability was further accentuated, primarily through increase in goal-equivalent variance. This provides evidence for the hypothesis that under challenging task constraints increased variability is selectively directed into task-irrelevant degrees of freedom. Because differential control along different directions of joint space requires coordination among joint angles, this observation supports the view that the CNS responds to increased task difficulty through enhanced coordination among degrees of freedom. The adaptive nature of this coordination is further illustrated by the similar enhanced use of goal-equivalent joint combinations to achieve a stable CM position when subjects stood up under the additional constraint of maintaining light touch with the fingertips. This was achieved by channeling goal-equivalent variability into different directions of joint configuration space.     

足部跖趾关节约束对人体步态稳定性的影响

目的 研究足部跖趾关节约束对人体步态稳定性的影响。 方法 在水平湿滑试验台上进行足部跖趾关节有、无约束两种状态下的步态实验,分析时空步态参数、运动学参数、动力学参数以及利用摩擦因数( utilization coefficient of friction,UCOF)差异。 结果 跖趾关节有约束状态下,人体行走平均步速减小 50 mm/ s,跨步长度缩小0. 22 m,双支撑相时间缩短 70 ms;跖趾关节约束会使髋、膝关节在矢状面内的活动范围显著增大,而踝关节活动范围减小。 同时,跖趾关节约束状态 UCOF 幅值是无约束状态的 1. 15 倍,表明人体滑跌的概率增大以及行走不稳定性增加。 结论 足部跖趾关节约束会降低行走稳定性。 研究结果为足部趾屈运动康复设备的研发提供数据和理论支持     

The role of active forces and intersegmental dynamics in the control of limb trajectory over obstacles during locomotion in humans

The focus of this paper is to examine the contributions of active and passive forces in the control of limb trajectory over obstacles during locomotion. Kintetic analyses of the swing phase of locomotion were carried out to determine the power profiles at various joints and to parcel the joint moments into moments due to muscle action, gravitational force and motion-dependent terms. The analyses revealed that toe elevation over the obstacles was achieved primarily by flexing at the hip, knee and ankle joint. Power analyses showed that translational energy applied at the hip joint and rotational energy applied at the knee joint were modulated as functions of obstacle height. This demonstrates that increased hip and ankle joint flexion are achieved not through active muscle action but rather through passive forces induced by translational action at the hip (representing contribution by the stance limb muscles) and rotational action at the knee joint. Parcelling the joint moment terms into various components clearly shows how the nervous system exploits intersegmental dynamics to simplify control of limb elevation over obstacles and minimize energy costs.     

Modulation of simple reaction time on the background of an oscillatory action: implications for synergy organization

The hypothesis put forward here is that simple reaction time (SRT) modulation on the background of an oscillatory motor action is due to central neural coupling between signals to the effectors involved in the focal and the oscillatory action. The strength of the coupling may be defined by various factors ranging from anatomy to personal lifetime practice or to a particular task context. In one experiment, subjects performed an SRT task (ipsi or contralateral elbow flexion or ipsilateral ankle plantar flexion) in response to a visual imperative signal presented during a continuous oscillatory movement of the right wrist. Discrete elbow movements lead to nearly simultaneous large bursts of activity in both biceps and the wrist flexor of the arm. Strong modulation of premotor time (peak to peak changes of about 80 ms) with the phase of oscillatory action (f(OSC)) was seen in both biceps and wrist flexor when the two movements were performed by joints of the same limb but not when they were performed by joints of different limbs. The order of recruitment of proximal and distal muscles was also dependent on the phase of oscillatory action: the typical proximal-to-distal order was seen at relatively long premotor times (PMTs) while simultaneous muscle activation was seen at the shortest PMTs. In the second experiment, the subjects held a cylindrical plastic cup in the left hand and applied sine-like isometric force to the bottom of the cup with the other hand. The SRT in the task requiring a quick increase in the grip force in response to a visual imperative stimulus was modulated with the phase of the oscillatory action. This modulation disappeared when the right hand applied similarly modulated force to another surface. The conclusion is that an interaction between control signals for the focal and oscillatory actions at a supraspinal level led to the observed modulation of the SRT during the phase of oscillatory action. The possible role of cortical and subcortical mechanisms is discussed.     

Ascending spinal axons that signal the position of the hindlimbs under static conditions: Location and receptor input

Summary Psychophysical experiments have shown that signals from slowly adapting subcutaneous receptors are used to sense limb position under static conditions (i.e., when the joints are stationary). The ascending collaterals of the slowly adapting primary sensory neurons supplying the deep tissues of the hindlimb do not project to the brain via the fasciculus gracilis. In experiments on cats, we have found a population of axons in the lateral fasciculus that signal the position of the ipsilateral hindlimb with a slowly adapting discharge. In the lower thoracic cord these fibers lie between the spinocervical tract and the ventral roots. Although plentiful in the lower thoracic cord, they are sparse or absent below L₃. In addition, a few position signaling axons with crossed input were found in the ventral part of the lateral white matter and in the ventral columns. Since the clinical evidence suggests that the spinal pathway for position sense is uncrossed, we propose that information used for conscious judgments of limb position when the joints are stationary initially ascends via the dorsal columns and then relays to the lateral fasciculus on the same side. These slowly adapting signals also may be used to judge limb position when the joints are moving. To determine whether this slowly adapting discharge originates from muscle or joint receptors, the tendons crossing the ankle joint were exposed but left in continuity and then pulled on while the joint was stationary. In this way individual lateral fascicular axons that signaled ankle flexion, extension, abduction or adduction could be shown to receive a strong excitatory input from muscle receptors. After the muscle tendons crossing the ankle joint were cut, tract fibers signaling ankle flexion, extension, abduction or adduction could no longer be found in this portion of the spinal white matter. Axons signaling clockwise or counterclock-wise twist of the ankle were reduced in number but a few were still present. These results suggest that muscle receptors provide the predominant signal used to sense ankle flexion, extension, abduction and adduction and that receptors in articular tissues may signal ankle twist.