Multijoint arm movements in cerebellar ataxia: Abnormal control of movement dynamics

In cerebellar ataxia, kinematic aberrations of multijoint movements are thought to originate from deficiencies in generating muscular torques that are adequate to control the mechanical consequences of dynamic interaction forces. At this point the exact mechanisms that lead to an abnormal control of interaction torques are not known. In principle, the generation of inadequate muscular torques may result from an impairment in generating sufficient levels of torques or from an inaccurate assessment and prediction of the mechanical consequences of movements of one limb segment on adjacent joints. We sought to differentiate the relative contribution of these two mechanisms and, therefore, analyzed intersegmental dynamics of multijoint pointing movements in healthy subjects and in patients with cerebellar degeneration. Unrestrained vertical arm movements were performed at three different target movement velocities and recorded using an optoelectronic tracking system. An inverse dynamics approach was employed to compute net joint torques, muscular torques, dynamic interaction torques and gravitational torques acting at the elbow and shoulder joint. In both groups, peak dynamic interaction forces and peak muscular forces were largest during fast movements. In contrast to normal subjects, patients produced hypermetric movements when executing fast movements. Hypermetric movements were associated with smaller peak muscular torques and smaller rates of torque change at elbow and shoulder joints. The patients’ deficit in generating appropriate levels of muscular force were prominent during two different phases of the pointing movement. Peak muscular forces at the elbow were reduced during the initial phase of the movement when simultaneous shoulder joint flexion generated an extensor influence upon the elbow joint. When attempting to terminate the movement, gravitational and dynamic interaction forces caused overshooting extension at the elbow joint. In normal subjects, muscular torque patterns at shoulder and elbow joint were synchronized in that peak flexor and extensor muscular torques occurred simultaneously at both joints. This temporal pattern of muscular torque generation at shoulder and elbow joint was preserved in patients. Our data suggest that an impairment in generating sufficient levels of phasic muscular torques significantly contributes to the patients’ difficulties in controlling the mechanical consequences of dynamic interaction forces during multijoint movements. Received: 28 October 1996 / Accepted: 30 September 1997     

Deliberate utilization of interaction torques brakes elbow extension in a fast throwing motion

We tested the hypothesis that in fast arm movements the CNS deliberately utilizes interaction torques to decelerate (brake) joint rotations. Twelve subjects performed fast 2-D overarm throws in which large elbow extension velocities occurred. Joint motions were computed from recordings made with search coils; joint torques were calculated using inverse dynamics. After ball release, a large follow-through shoulder extension acceleration occurred that was initiated by shoulder extensor muscle torque. This shoulder acceleration produced a flexor interaction torque at the elbow that initiated elbow deceleration (braking). An instantaneous mechanical interaction of passive torques then occurred between elbow and shoulder, i.e., elbow extension deceleration produced a large shoulder extensor interaction torque that contributed to the shoulder extension acceleration which, simultaneously, produced a large elbow flexor interaction torque that contributed to elbow extension deceleration, and so on. Late elbow flexor muscle torque also contributed to elbow deceleration. The interaction of passive torques between shoulder and elbow was braked by shoulder flexor muscle torque. In this mechanism, shoulder musculature contributed to braking elbow extension in two ways: shoulder extensors initiated the mechanical interaction of passive torques between shoulder and elbow and shoulder flexors dissipated kinetic energy from elbow braking. It is concluded that, in fast 2-D throws, the CNS deliberately utilizes powerful interaction torques between shoulder and elbow to brake motion at the elbow.     

Independent coactivation of shoulder and elbow muscles

 The aim of this study was to examine the possibility of independent muscle coactivation at the shoulder and elbow. Subjects performed rapid point-to-point movements in a horizontal plane from different initial limb configurations to a single target. EMG activity was measured from flexor and extensor muscles acting at the shoulder (pectoralis clavicular head and posterior deltoid) and elbow (biceps long head and triceps lateral head) and flexor and extensor muscles acting at both joints (biceps short head and triceps long head). Muscle coactivation was assessed by measuring tonic levels of electromyographic (EMG) activity after limb position stabilized following the end of the movements. It was observed that tonic EMG levels following movements to the same target varied as a function of the amplitude of shoulder and elbow motion. Moreover, for the movements tested here, the coactivation of shoulder and elbow muscles was found to be independent – tonic EMG activity of shoulder muscles increased in proportion to shoulder movement, but was unrelated to elbow motion, whereas elbow and double-joint muscle coactivation varied with the amplitude of elbow movement and were not correlated with shoulder motion. In addition, tonic EMG levels were higher for movements in which the shoulder and elbow rotated in the same direction than for those in which the joints rotated in opposite directions. In this respect, muscle coactivation may reflect a simple strategy to compensate for forces introduced by multijoint limb dynamics. Received: 7 July 1998 / Accepted: 28 July 1998     

A novel shoulder–elbow mechanism for increasing speed in a multijoint arm movement

The speed of arm movements is normally increased by increasing agonist muscle activity, but in overarm throwing, an additional effect on speed may come from exploitation of interaction torques (a passive torque associated with motion at adjacent joints). We investigated how the central nervous system (CNS) controls interaction torques at the shoulder and elbow to increase speed in 2-D overarm throwing. Twelve experienced throwers made slow, medium, and fast 2-D throws in a parasagittal plane. Joint motions were computed from recordings made with search coils; joint torques were calculated using inverse dynamics. For slow and medium-speed throws, elbow extension was primarily produced by elbow muscle torque. For fast throws, there was an additional late-occurring elbow extensor interaction torque. Parceling out this elbow extension interaction torque revealed that it primarily arose from shoulder extension deceleration. Surprisingly, shoulder deceleration before ball release was not caused by shoulder flexor (antagonist) muscle torque. Rather, shoulder deceleration was produced by passive elbow-to-shoulder interaction torques that were primarily associated with elbow extension acceleration and velocity. It is concluded that when generating fast 2-D throws, the CNS utilized the arm’s biomechanical properties to increase ball speed. It did this by coordinating shoulder and elbow motions such that an instantaneous mechanical positive feedback occurred of interaction torques between shoulder and elbow before ball release. To what extent this mechanism is utilized in other fast multijoint arm movements remains to be determined.     

Effects of inactivation of the anterior interpositus nucleus on the kinematic and dynamic control of multijoint movement

We previously showed that inactivating the anterior interpositus nucleus in cats disrupts prehension; paw paths, normally straight and accurate, become curved, hypometric, and more variable. In the present study, we determined the joint kinematic and dynamic origins of this impairment. Animals were restrained in a hammock and trained to reach and grasp a cube of meat from a narrow food well at varied heights; movements were monitored using the MacReflex analysis system. The anterior interpositus nucleus was inactivated by microinjection of the GABA agonist muscimol (0.25-0.5 microgram in 0.5 microliter saline). For each joint, we computed the torque due to gravity, inertial resistance (termed self torque), interjoint interactions (termed interaction torque), and the combined effects of active muscle contraction and passive soft tissue stretch (termed generalized muscle torque). Inactivation produced significant reductions in the amplitude, velocity, and acceleration of elbow flexion. However, these movements continued to scale normally with target height. Shoulder extension was reduced by inactivation but wrist angular displacement and velocity were not. Inactivation also produced changes in the temporal coordination between elbow, shoulder, and wrist kinematics. Dynamic analysis showed that elbow flexion both before and during inactivation was produced by the combined action of muscle and interaction torque, but that the timing depended on muscle torque. Elbow interaction and muscle torques were scaled to target height both before and during inactivation. Inactivation produced significant reductions in elbow flexor interaction and muscle torques. The duration of elbow flexor muscle torque was prolonged to compensate for the reduction in flexor interaction torque. Shoulder extension was produced by extensor interaction and muscle torques both before and during inactivation. Inactivation produced a reduction in shoulder extension, primarily by reduced interaction torque, but without compensation. Wrist plantarflexion, which occurred during elbow flexion, was driven by plantarflexor interaction and gravitational torques both before and during inactivation. Muscle torque acted in the opposite direction with a phase lead to restrain the plantarflexor interaction torque. During inactivation, there was a reduction in plantarflexor interaction torque and a loss of the phase lead of the muscle torque. Our findings implicate the C1/C3 anterior interpositus zone of the cerebellum in the anticipatory control of intersegmental dynamics during reaching, which zone is required for coordinating the motions of the shoulder and wrist with those of the elbow. In contrast, this cerebellar zone does not play a role in scaling the movement to match a target.     

The development of goal-directed reaching in infants: hand trajectory formation and joint torque control

Nine young infants were followed longitudinally from 4 to 15 months of age. We recorded early spontaneous movements and reaching movements to a stationary target. Time-position data of the hand (endpoint), shoulder, and elbow were collected using an optoelectronic measurement system (ELITE). We analyzed the endpoint kinematics and the intersegmental dynamics of the shoulder and elbow joint to investigate how changes in proximal torque control determined the development of hand trajectory formation. Two developmental phases of hand trajectory formation were identified: a first phase of rapid improvements between 16 and 24 weeks of age, the time of reaching onset for all infants. During that time period the number of movement units per reach and movement time decreased dramatically. In a second phase (28–64 weeks), a period of fine-tuning of the sensorimotor system, we saw slower, more gradual changes in the endpoint kinematics. The analysis of the underlying intersegmental joint torques revealed the following results: first, the range of muscular and motiondependent torques (relative to body weight) did not change significantly with age. That is, early reaching was not confined by limitations in producing task-adequate levels of muscular torque. Second, improvements in the endpoint kinematics were not accomplished by minimizing amplitude of muscle and reactive torques. Third, the relative timing of muscular and motion-dependent torque peaks showed a systematic development toward an adult timing profile with increasing age. In conclusion, the development toward invariant characteristics of the hand trajectory is mirrored by concurrent changes in the control of joint forces. The acquisition of stable patterns of intersegmental coordination is not achieved by simply regulating force amplitude, but more so by modulating the correct timing of joint force production and by the system's use of reactive forces. Our findings support the view that development of reaching is a process of unsupervised learning with no external or innate teacher prescribing the desired kinematics or kinetics of the movement.     

Kinematics and kinetics of multijoint reaching in nonhuman primates

The present study identifies the mechanics of planar reaching movements performed by monkeys (Macaca mulatta) wearing a robotic exoskeleton. This device maintained the limb in the horizontal plane such that hand motion was generated only by flexor and extensor motions at the shoulder and elbow. The study describes the kinematic and kinetic features of the shoulder, elbow, and hand during reaching movements from a central target to peripheral targets located on the circumference of a circle: the center-out task. While subjects made reaching movements with relatively straight smooth hand paths and little variation in peak hand velocity, there were large variations in joint motion, torque, and power for movements in different spatial directions. Unlike single-joint movements, joint kinematics and kinetics were not tightly coupled for these multijoint movements. For most movements, power generation was predominantly generated at only one of the two joints. The present analysis illustrates the complexities inherent in multijoint movements and forms the basis for understanding strategies used by the motor system to control reaching movements and for interpreting the response of neurons in different brain regions during this task.     

Effect of sensory feedback from the proximal upper limb on voluntary isometric finger flexion and extension in hemiparetic stroke subjects

This study investigated the potential influence of proximal sensory feedback on voluntary distal motor activity in the paretic upper limb of hemiparetic stroke survivors and the potential effect of voluntary distal motor activity on proximal muscle activity. Ten stroke subjects and 10 neurologically intact control subjects performed maximum voluntary isometric flexion and extension, respectively, at the metacarpophalangeal (MCP) joints of the fingers in two static arm postures and under three conditions of electrical stimulation of the arm. The tasks were quantified in terms of maximum MCP torque [MCP flexion (MCP(flex)) or MCP extension (MCP(ext))] and activity of targeted (flexor digitorum superficialis or extensor digitorum communis) and nontargeted upper limb muscles. From a previous study on the MCP stretch reflex poststroke, we expected stroke subjects to exhibit a modulation of voluntary MCP torque production by arm posture and electrical stimulation and increased nontargeted muscle activity. Posture 1 (flexed elbow, neutral shoulder) led to greater MCP(flex) in stroke subjects than posture 2 (extended elbow, flexed shoulder). Electrical stimulation did not influence MCP(flex) or MCP(ext) in either subject group. In stroke subjects, posture 1 led to greater nontargeted upper limb flexor activity during MCP(flex) and to greater elbow flexor and extensor activity during MCP(ext). Stroke subjects exhibited greater elbow flexor activity during MCP(flex) and greater elbow flexor and extensor activity during MCP(ext) than control subjects. The results suggest that static arm posture can modulate voluntary distal motor activity and accompanying muscle activity in the paretic upper limb poststroke.     

Novel muscle patterns for reaching after cervical spinal cord injury: a case for motor redundancy

A fundamental issue in the neuromotor control of arm movements is whether the nervous system can use distinctly different muscle activity patterns to obtain similar kinematic outcomes. Although computer simulations have demonstrated several possible mechanical and torque solutions, there is little empirical evidence that the nervous system actually employs fundamentally different muscle patterns for the same movement, such as activating a muscle one time and not the next, or switching from a flexor to an extensor. Under typical conditions, subjects choose the same muscles for any given movement, which suggests that in order to see the capacity of the nervous system to make a different choice of muscles, the nervous system must be pushed beyond the normal circumstances. The purpose of this study, then, was to examine an atypical condition, reaching of cervical spinal cord injured (SCI) subjects who have a reduced repertoire of available distal arm muscles but otherwise a normal nervous system above the level of lesion. Electromyography and kinematics of the shoulder and elbow were examined in the SCI subjects performing a center-out task and then compared to neurologically normal control subjects. The findings showed that the SCI-injured subjects produced reaches with typical global kinematic features, such as straight finger paths, bell-shaped velocities, and joint excursions similar to control subjects. The SCI subjects, however, activated only the shoulder agonist muscle for all directions, unlike the control pattern that involved a reciprocal pattern at each joint (shoulder, elbow, and wrist). Nonetheless, the SCI subjects could activate their shoulder antagonist muscles, elbow flexors, and wrist extensor (extensor carpi radialis) for isometric tasks, but did not activate them during the reaching movements. These results demonstrate that for reaching movements, the SCI subjects used a strikingly different pattern of intact muscle activities than control subjects. Hence, the findings imply that the nervous system is capable of choosing either the control pattern or the SCI pattern. We would speculate that control subjects do not select the SCI pattern because the different choice of muscles results in kinematic features (reduced fingertip speed, multiple shoulder accelerations) other than the global features that are somehow less advantageous or efficient.     

Inter-joint coupling strategy during adaptation to novel viscous loads in human arm movement

When arm movements are perturbed by a load, how does the nervous system adjust control signals to reduce error? While it has been shown that the nervous system is capable of compensating for the effects of limb dynamics and external forces, the strategies used to adapt to novel loads are not well understood. We used a robotic exoskeleton [kinesiological instrument for normal and altered reaching movements (KINARM)] to apply novel loads to the arm during single-joint elbow flexions in the horizontal plane (shoulder rotation was allowed). Loads varied in magnitude with the instantaneous velocity of elbow flexion, and were applied to the shoulder in experiment 1 (interaction loads) and the elbow in experiment 2 (direct loads). Initial exposure to both interaction and direct loads resulted in perturbations at both joints, even though the load was applied to only a single joint. Subjects tended to correct for the kinematics of the elbow joint while perturbations at the shoulder persisted. Electromyograms (EMGs) and computed muscle torque showed that subjects modified muscle activity at the elbow to reduce elbow positional deviations. Shoulder muscle activity was also modified; however, these changes were always in the same direction as those at the elbow. Current models of motor control based on inverse-dynamics calculations and force-control, as well as models based on positional control, predict an uncoupling of shoulder and elbow muscle torques for adaptation to these loads. In contrast, subjects in this study adopted a simple strategy of modulating the natural coupling that exists between elbow and shoulder muscle torque during single-joint elbow movements.     

Cerebellar ataxia: torque deficiency or torque mismatch between joints?

Prior work has shown that cerebellar subjects have difficulty adjusting for interaction torques that occur during multi-jointed movements. The purpose of this study was to determine whether this deficit is due to a general inability to generate sufficient levels of phasic torque inability or due to an inability to generate muscle torques that predict and compensate for interaction torques. A second purpose was to determine whether reducing the number of moving joints by external mechanical fixation could improve cerebellar subjects' targeted limb movements. We studied control and cerebellar subjects making elbow flexion movements to touch a target under two conditions: 1) a shoulder free condition, which required only elbow flexion, although the shoulder joint was unconstrained and 2) a shoulder fixed condition, where the shoulder joint was mechanically stabilized so it could not move. We measured joint positions of the arm in the sagittal plane and electromyograms (EMGs) of shoulder and elbow muscles. Elbow and shoulder torques were estimated using inverse dynamics equations. In the shoulder free condition, cerebellar subjects made greater endpoint errors (primarily overshoots) than did controls. Cerebellar subjects' overshoot errors were largely due to unwanted flexion at the shoulder. The excessive shoulder flexion resulted from a torque mismatch, where larger shoulder muscle torques were produced at higher rates than would be appropriate for a given elbow movement. In the shoulder fixed condition, endpoint errors of cerebellar subjects and controls were comparable. The improved accuracy of cerebellar subjects was accompanied by reduced shoulder flexor muscle activity. Most of the correct cerebellar trials in the shoulder fixed condition were movements made using only muscles that flex the elbow. Our findings suggest that cerebellar subjects' poor shoulder control is due to an inability to generate muscle torques that predict and compensate for interaction torques, and not due to a general inability to generate sufficient levels of phasic torque. In addition, reducing the number of muscles to be controlled improved cerebellar ataxia.     

The leading joint hypothesis for spatial reaching arm motions

The leading joint hypothesis (LJH), developed for planar arm reaching, proposes that the interaction torques experienced by the proximal joint are low compared to the corresponding muscle torques. The human central nervous system could potentially ignore these interaction torques at the proximal (leading) joint with little effect on the wrist trajectory, simplifying joint-level control. This paper investigates the extension of the LJH to spatial reaching. In spatial motion, a number of terms in the governing equation (Euler’s angular momentum balance) that vanish for planar movements are non-trivial, so their contributions to the joint torque must be classified as net, interaction or muscle torque. This paper applies definitions from the literature to these torque components to establish a general classification for all terms in Euler’s equation. This classification is equally applicable to planar and spatial motion. Additionally, a rationale for excluding gravity torques from the torque analysis is provided. Subjects performed point-to-point reaching movements between targets whose locations ensured that the wrist paths lay in various portions of the arm’s spatial workspace. Movement kinematics were recorded using electromagnetic sensors located on the subject’s arm segments and thorax. The arm was modeled as a three-link kinematic chain with idealized spherical and revolute joints at the shoulder and elbow. Joint torque components were computed using inverse dynamics. Most movements were ‘shoulder-led’ in that the interaction torque impulse was significantly lower than the muscle torque impulse for the shoulder, but not the elbow. For the few elbow-led movements, the interaction impulse at the elbow was low, while that at the shoulder was high, and these typically involved large elbow and small shoulder displacements. These results support the LJH and extend it to spatial reaching motion.     

General coordination of shoulder,elbow and wrist dynamics during multijoint arm movements

Studies of multijoint arm movements have demonstrated that the nervous system anticipates and plans for the mechanical effects that arise from motion of the linked limb segments. The general rules by which the nervous system selects appropriate muscle activities and torques to best deal with these intersegmental effects are largely unknown. In order to reveal possible rules, this study examined the relationship of muscle and interaction torques to joint acceleration at the shoulder, elbow and wrist during point-to-point arm movements to a range of targets in the horizontal plane. Results showed that, in general, dynamics differed between the joints. For most movements, shoulder muscle torque primarily determined net torque and joint acceleration, while interaction torque was minimal. In contrast, elbow and wrist net torque were determined by a combination of muscle and interaction torque that varied systematically with target direction and joint excursion. This "shoulder-centered pattern" occurred whether subjects reached targets using straight or curved finger paths. The prevalence of a shoulder-centered pattern extends findings from a range of arm movement studies including movement of healthy adults, neurological patients, and simulations with altered interaction effects. The shoulder-centered pattern occurred for most but not all movements. The majority of the remaining movements displayed an "elbow-centered pattern," in which muscle torque determined initial acceleration at the elbow and not at the shoulder. This occurred for movements when shoulder excursion was <50% of elbow excursion. Thus, both shoulder- and elbow-centered movements displayed a difference between joints but with reversed dynamics. Overall, these findings suggest that a difference in dynamics between joints is a general feature of horizontal plane arm movements, and this difference is most commonly reflected in a shoulder-centered pattern. This feature fits well with other general shoulder-elbow differences suggested in the literature on arm movements, namely that: (a) agonist muscle activity appears more closely related to certain joint kinematics at the shoulder than at the elbow, (b) adults with neurological damage display less disruption of shoulder motion than elbow motion, and (c) infants display adult-like motion first in the shoulder and last at the wrist.     

The influence of agonist premotor silence and the stretch-shortening cycle on contractile rate in active skeletal muscle

Summary Agonist premotor silence (PMS), a brief period of relative quiescence in active skeletal muscle prior to phasic activation, was investigated in subjects performing maximal contractions. The frequency of occurrence and potential function of the silent period were examined for elbow flexions and extensions. PMS was evident for movements in both directions, indicating that the mechanism is not primarily limited to extensors as previously hypothesized. Flexions demonstrating PMS exhibited increased velocity and acceleration; however, kinematic facilitation was only evident on trials exhibiting the muscular stretch-shortening cycle (SSC). The SSC was present on trials lacking PMS, demonstrating that biceps and triceps silence are not the sole determinants of preparatory agonist lengthening for elbow flexions and extensions, respectively. Taken together, the data indicate that agonist PMS is a mechanism under apparent central control that acts concomitantly with mechanical factors to potentiate elbow flexor contractions.     

Sensory-motor equivalence: manual aiming in C6 tetraplegics following musculotendinous transfer surgery at the elbow

Cervical spinal lesions at C6 result in paralysis of the triceps brachii while leaving deltoid and elbow flexor function intact. We examined the spatial–temporal characteristics of goal-directed aiming movements performed by C6 tetraplegics who had undergone musculotendinous transfer surgery in which the posterior deltoid replaces the triceps as the elbow extensor. On some trials, liquid crystal goggles were used to eliminate vision of the limb and target upon movement initiation. Although tetraplegic participants achieved the same degree of movement accuracy/consistency as control participants, their movement times were longer regardless of whether the movements were made away from (elbow extension) or towards the body (elbow flexion). Longer movement times were related to lower peak velocities, and not the symmetry of the aiming profiles. The tetraplegic participants were no more dependent on visual feedback for limb regulation than control participants. Although the characteristics of the movement trajectories were surprisingly similar, in both vision conditions, tetraplegics required more real and proportional time to reduce spatial variability in the limb’s trajectory for elbow extensions. Our results indicate that the sensorimotor system is adaptable and that the representations governing limb control are not muscle specific.     

Compensation for interaction torques during single- and multijoint limb movement

During multijoint limb movements such as reaching, rotational forces arise at one joint due to the motions of limb segments about other joints. We report the results of three experiments in which we assessed the extent to which control signals to muscles are adjusted to counteract these "interaction torques." Human subjects performed single- and multijoint pointing movements involving shoulder and elbow motion, and movement parameters related to the magnitude and direction of interaction torques were manipulated systematically. We examined electromyographic (EMG) activity of shoulder and elbow muscles and, specifically, the relationship between EMG activity and joint interaction torque. A first set of experiments examined single-joint movements. During both single-joint elbow (experiment 1) and shoulder (experiment 2) movements, phasic EMG activity was observed in muscles spanning the stationary joint (shoulder muscles in experiment 1 and elbow muscles in experiment 2). This muscle activity preceded movement and varied in amplitude with the magnitude of upcoming interaction torque (the load resulting from motion of the nonstationary limb segment). In a third experiment, subjects performed multijoint movements involving simultaneous motion at the shoulder and elbow. Movement amplitude and velocity at one joint were held constant, while the direction of movement about the other joint was varied. When the direction of elbow motion was varied (flexion vs. extension) and shoulder kinematics were held constant, EMG activity in shoulder muscles varied depending on the direction of elbow motion (and hence the sign of the interaction torque arising at the shoulder). Similarly, EMG activity in elbow muscles varied depending on the direction of shoulder motion for movements in which elbow kinematics were held constant. The results from all three experiments support the idea that central control signals to muscles are adjusted, in a predictive manner, to compensate for interaction torques-loads arising at one joint that depend on motion about other joints.     

Passive resting state and history of antagonist muscle activity shape active extensions in an insect limb

Limb movements can be driven by muscle contractions, external forces, or intrinsic passive forces. For lightweight limbs like those of insects or small vertebrates, passive forces can be large enough to overcome the effects of gravity and may even generate limb movements in the absence of active muscle contractions. Understanding the sources and actions of such forces is therefore important in understanding motor control. We describe passive properties of the femur-tibia joint of the locust hind leg. The resting angle is determined primarily by passive properties of the relatively large extensor tibiae muscle and is influenced by the history of activation of the fast extensor tibiae motor neuron. The resting angle is therefore better described as a history-dependent resting state. We selectively stimulated different flexor tibiae motor neurons to generate a range of isometric contractions of the flexor tibiae muscle and then stimulated the fast extensor tibiae motor neuron to elicit active tibial extensions. Residual forces in the flexor muscle have only a small effect on subsequent active extensions, but the effect is larger for distal than for proximal flexor motor neurons and varies with the strength of flexor activation. We conclude that passive properties of a lightweight limb make substantial and complex contributions to the resting state of the limb that must be taken into account in the patterning of neuronal control signals driving its active movements. Low variability in the effects of the passive forces may permit the nervous system to accurately predict their contributions to behavior.     

等动肌力矩的增龄性发育特征

背景：少年时期肌肉力量是否能够得到平衡发育对其生长发育过程中起重要作用。 目的：探索少年的肌力发展的生物力学规律。 方法：利用Kinitech等速测力系统对中小学男女生各30名双侧肩、髋关节屈伸肌群进行等动肌力测试,测试速度为60 (°)/s,测试次数为屈伸6次。 结果与结论：所有测试者的髋、肩关节屈伸肌的相对峰力矩随测试速度的增大而逐渐减小;且随年龄增长肩、髋关节屈伸肌的相对峰力矩均同步增大;小学生在不同测试速度状态下男生髋、肩关节相对峰力矩均大于女生(P < 0.05),但中学生髋、肩关节相对峰力矩没有性别差异。     

Wrist muscle activation,interaction torque and mechanical properties in unskilled throws of different speeds

An unexpected property of unskilled overarm throws is that wrist flexion velocity at ball release does not increase in throws of increasing speed. We investigated the nature of the interaction torques and wrist mechanical properties that have been proposed to produce this property. Twelve recreational throwers made seated 2-D throws, which were used as a model for unskilled throwing. Joint motions were computed from recordings made with search coils; joint torques were calculated from inverse dynamics. Wrist flexion velocity at ball release was actually smaller in fast throws than in slow throws. This was associated in fast throws with the decrease in a large wrist flexor muscle torque (i.e., a calculated residual torque) in the last 40 ms before ball release, and its reversal to an extensor torque. Consequently, wrist flexor muscle torque was unable to oppose a small maintained wrist extensor interaction torque that arose from continuing elbow extension acceleration. The decrease in wrist flexor muscle torque was not associated with a decrease in wrist flexor EMG activity, nor with an increase in wrist extensor EMG activity. These findings support the hypothesis that the smaller wrist flexion velocity at ball release in fast 2-D throws results from a wrist extensor interaction torque and from a large wrist extensor viscoelastic torque. We propose that in fast 3-D throws skilled subjects decelerate elbow extension before ball release to help overcome these wrist extensor torques.     

Nonuniform distribution of reach-related and torque-related activity in upper arm muscles and neurons of primary motor cortex

The present study examined the activity of primate shoulder and elbow muscles using a novel reaching task. We enforced similar patterns of center-out movement while the animals countered viscous loads at their shoulder, elbow, both joints, or neither joint. Accordingly, we could examine reach-related activity during the unloaded condition and torque-related activity by comparing activity across load conditions. During unloaded reaching the upper arm muscles exhibited a bimodal distribution of preferred hand direction. Maximal reach-related activity occurred with hand movements mostly toward or away from the body. Arm muscles also exhibited a bimodal distribution of their preferred torque direction. Maximal torque-related activity typically occurred with shoulder-extension/elbow-flexion torque or shoulder-flexion/elbow-extension torque. Similar biases in reach-related and torque-related activity could be reproduced by optimizing a global measure of muscle activity. These biases were also observed in the neural activity of primary motor cortex (M1). The parallels between M1 and muscular activity demonstrate another link between motor cortical processing and the motor periphery and may reflect an optimization process performed by the sensorimotor system.