St-Onge N, Adamovich S V, Feldman A G
Department of Physiology, University of Montreal, Quebec, Canada.
Neuroscience. 1997 Jul;79(1):295-316. doi: 10.1016/s0306-4522(97)00071-7.
Using a non-linear dynamic model based on the lambda version of the equilibrium-point hypothesis, we investigated the shape and duration of the control patterns underlying discrete elbow movements. The model incorporates neural control variables, time-, position- and velocity-dependent intrinsic muscle and reflex properties. Two control variables (R and C) specify a positional frame of reference for activation of flexor and extensor motoneurons. The variable R (reciprocal command) specifies the referent joint angle (R) at which the transition of net flexor to extensor active torque or vice versa can be observed during changes in the actual joint angle elicited by an external force. The variable C (coactivation command) surrounds the transition angle by an angular range in which flexor and extensor muscles may be simultaneously active (if C > 0) or silent (if C < or = 0). An additional, time-dimensional control variable (mu command) influences the dependency of the threshold of the stretch reflex on movement velocity. This control variable is responsible for the reflex damping. Changes in the R command result in shifts in the equilibrium state of the system, a dynamical process leading to electromyographic modifications and movement production. Commands C and mu provide movement stability and effective energy dissipation preventing oscillations at the end of movement. A comparison of empirical and model data was carried out. A monotonic ramp-shaped pattern of the R command can account for the empirical kinematic and electromyographic patterns of the fastest elbow flexion movements made with or without additional inertia, as well as of self-paced movements. The rate of the shifts used in simulation was different for the three types of movements but independent of movement distance (20-80 degrees). This implies that, for a given type of movement, the distance is encoded by the duration of shift in the equilibrium state. The model also reproduces the kinematic and electromyographic patterns of the fastest uncorrected movements opposed in random trials by a high load (80-90% of the maximal) generated by position feedback to a torque motor. The following perturbation effects were simulated: a substantial decrease in the arm displacement (from 60-70 degrees to 5-15 degrees) and movement duration (to about 100 ms) so that these movements ended near the peak velocity of those which were not perturbed; a prolongation of the first agonist electromyographic burst as long as the load was applied; the suppression of the antagonist burst during the dynamic and static phases of movements: the reappearance of the antagonist burst in response to unloading accompanied by a short-latency suppression of agonist activity. These kinematic and electromyographic features of both perturbed and non-perturbed movements were reproduced by using the same control patterns which elicited a monotonic shift in the equilibrium state of the system ending before the peak velocity of non-perturbed movements. Our results suggest that the neural control processes underlying the fastest unopposed changes in the arm position are completed long before the end of the movement and phasic electromyographic activity. Neither the timing nor the amplitude of electromyographic bursts are planned but rather they represent the long-lasting dynamic response of central, reflex and mechanical components of the system to a monotonic, short-duration shift in the system's equilibrium state.
基于平衡点假设的拉姆达版本,我们使用非线性动力学模型研究了离散肘部运动背后控制模式的形状和持续时间。该模型纳入了神经控制变量、与时间、位置和速度相关的内在肌肉和反射特性。两个控制变量(R和C)为屈肌和伸肌运动神经元的激活指定了一个位置参考系。变量R(反向指令)指定了参考关节角度(R),在由外力引起的实际关节角度变化期间,可以观察到净屈肌到伸肌主动扭矩的转变,反之亦然。变量C(共同激活指令)通过一个角度范围围绕转变角度,在该范围内屈肌和伸肌可能同时激活(如果C>0)或静止(如果C≤0)。另一个时间维度的控制变量(μ指令)影响牵张反射阈值对运动速度的依赖性。这个控制变量负责反射阻尼。R指令的变化导致系统平衡状态的转变,这是一个导致肌电图改变和运动产生的动态过程。指令C和μ提供运动稳定性和有效的能量耗散,防止运动结束时的振荡。对经验数据和模型数据进行了比较。R指令的单调斜坡形模式可以解释在有或没有额外惯性的情况下以及自定步速运动中最快肘部屈曲运动的经验运动学和肌电图模式。模拟中使用的转变速率对于三种运动类型是不同的,但与运动距离(20 - 80度)无关。这意味着,对于给定类型的运动,距离由平衡状态转变的持续时间编码。该模型还再现了在随机试验中由位置反馈到扭矩电机产生的高负载(最大负载的80 - 90%)对抗的最快未校正运动的运动学和肌电图模式。模拟了以下扰动效应:手臂位移大幅减少(从60 - 70度到5 - 15度)和运动持续时间缩短(至约100毫秒),以使这些运动在未受扰动运动的峰值速度附近结束;只要施加负载,第一个主动肌肌电图爆发就会延长;在运动的动态和静态阶段拮抗肌爆发受到抑制;卸载时拮抗肌爆发重新出现,同时伴随主动肌活动的短潜伏期抑制。通过使用相同的控制模式再现了受扰动和未受扰动运动的这些运动学和肌电图特征,这些控制模式在系统平衡状态产生单调转变,在未受扰动运动的峰值速度之前结束。我们的结果表明,手臂位置最快的无对抗变化背后的神经控制过程在运动结束和阶段性肌电图活动之前很久就已完成。肌电图爆发的时间和幅度都不是预先计划好的,而是代表了系统的中枢、反射和机械组件对系统平衡状态的单调、短持续时间转变的长期动态响应。