De Serres Sophie J, Bennett David J, Stein Richard B
Department of Physiology, University of Alberta, Edmonton, Alberta, Canada T6G 2H7.
J Physiol. 2002 Dec 15;545(3):1027-40. doi: 10.1113/jphysiol.2002.027177.
The triceps surae (TS) stretch reflex was measured in decerebrate cats during crossed extensor stimulation (tonic contractions) and after spinalization during rhythmic locomotor activity. The TS reflex force in response to a short pulse stretch measured during tonic contractions at low level of background activity was greater than when more background activity was present at the time of application of stretch. In contrast, the reflex force measured during rhythmic contractions was very small at low level of background force (flexion phase) and increased at moderate and high levels of background activity (extension phase). Thus, even in reduced preparations, a task modulation of the stretch reflex occurs. Throughout the experimental procedure, the torque motor used to stretch the muscles behaved like a spring of a preset compliance (from isometric to very compliant). A reflex model was used to simulate the responses obtained experimentally. The gain of the stretch reflex loop was estimated for each load condition and both behavioural tasks. The reflex loop gain was significantly larger as the compliance of the external load increased for both tonic and rhythmic contractions, although to a lesser extent in the phasically activated muscles. During rhythmic locomotor contractions the gain was less than 1, assuring stability of the system. In contrast, during tonic contractions against a compliant load the gain exceeded 1, consistent with the instability (oscillations, clonus) seen at times under these load conditions. However, the high gain and instability was only transient, since repeated stretch reduced the gain. Thus, non-linearities in the system assured vigorous responses at the onset of perturbations, but then weaker responses to ongoing perturbations to reduce the chance of feedback instability (clonus).
在交叉伸肌刺激(强直性收缩)期间,对去大脑猫的小腿三头肌(TS)牵张反射进行了测量,并在脊髓横断后对其进行了节律性运动活动测量。在低水平背景活动的强直性收缩期间,对短脉冲牵张的反应中,TS反射力大于牵张时存在更多背景活动时的反射力。相反,在节律性收缩期间测量的反射力在低水平背景力(屈曲阶段)时非常小,而在中等和高水平背景活动(伸展阶段)时增加。因此,即使在简化的实验准备中,牵张反射也会出现任务调制。在整个实验过程中,用于拉伸肌肉的扭矩电机表现得像一个具有预设顺应性(从等长到非常顺应)的弹簧。使用反射模型来模拟实验获得的反应。针对每种负荷条件和两种行为任务,估计了牵张反射环的增益。对于强直性和节律性收缩,随着外部负荷顺应性的增加,反射环增益显著增大,尽管在相位激活的肌肉中程度较小。在节律性运动收缩期间,增益小于1,确保了系统的稳定性。相反,在对抗顺应性负荷的强直性收缩期间,增益超过1,这与在这些负荷条件下有时出现的不稳定性(振荡、阵挛)一致。然而,高增益和不稳定性只是短暂的,因为重复牵张会降低增益。因此,系统中的非线性确保了在扰动开始时的强烈反应,但随后对持续扰动的反应较弱,以减少反馈不稳定性(阵挛)的可能性。