Duysens J, Pearson K G
Brain Res. 1980 Apr 14;187(2):321-32. doi: 10.1016/0006-8993(80)90206-1.
The role of proprioceptive input from the ankle extensor triceps surae in the control of walking was examined in premammillary cats walking on a treadmill. The left hindlimb was rigidly fixed in one position after denervating almost all the leg muscles except the ankle extensor (triceps surae) and ankle flexor (tibialis anterior). Rhythmic alternating contractions of the isolated ankle flexor and extensor occurred in the fixed hindleg during periods of walking in the other three intact limbs. These rhythmic contractions disappeared when the isolated triceps surae was stretched so as to increase the force of the active contractions to beyond 4 kg. With maintained stretch the periodic contractions in the ankle flexor and extensor returned only after the force in the stretched triceps surae gradually decreased and fell below approximately 4 kg. Isometric contractions of the triceps surae produced either by stimulation of ventral root S1 or by large amplitude vibrations also led to the sudden disappearance of ankle flexor bursts. Inhibition of the locomotory rhythm could also be produced in all muscles of a single intact hindleg by clamping the ankle joint in a flexed position so as to stretch the ankle extensor. In all these cases, an increased rate of stepping of the contralateral hindleg was associated with the inhibition of the rhythmic locomotory activity. It is suggested that triceps surae proprioceptors signalling the presence of loading of the hindlimb extensor muscles inhibit the central generation of hindlimb flexion. During normal walking this mechanism could be of major importance during stance to prevent the initiation of the swing phase of a time when hindlimb extension is fully needed to support the weight of the animal. Thus a necessary, but not always sufficient, condition for the initiation of swing may be an unloading of leg extensor muscles.
在跑步机上行走的乳头前猫中,研究了来自踝伸肌腓肠三头肌的本体感觉输入在行走控制中的作用。在使除踝伸肌(腓肠三头肌)和踝屈肌(胫骨前肌)外的几乎所有腿部肌肉去神经支配后,将左后肢刚性固定在一个位置。在其他三条完整肢体行走期间,孤立的踝屈肌和伸肌在固定的后肢中出现有节奏的交替收缩。当孤立的腓肠三头肌被拉伸以将主动收缩的力量增加到超过4千克时,这些有节奏的收缩消失。在持续拉伸的情况下,只有在被拉伸的腓肠三头肌中的力量逐渐减小并降至约4千克以下后,踝屈肌和伸肌中的周期性收缩才会恢复。通过刺激腹根S1或通过大幅度振动产生的腓肠三头肌等长收缩也导致踝屈肌爆发突然消失。通过将踝关节夹在屈曲位置以拉伸踝伸肌,也可以在单个完整后肢的所有肌肉中产生运动节律的抑制。在所有这些情况下,对侧后肢步幅增加与有节奏的运动活动抑制相关。有人认为,腓肠三头肌本体感受器发出后肢伸肌肌肉负荷存在的信号,抑制后肢屈曲的中枢产生。在正常行走过程中,这种机制在站立期间可能对防止在动物完全需要后肢伸展来支撑体重时启动摆动期具有重要意义。因此,启动摆动的一个必要但不总是充分的条件可能是腿部伸肌肌肉的卸载。