Allum J H, Honegger F
Department of ORL, University HNO-Klinik, Basel, Switzerland.
Exp Brain Res. 1998 Aug;121(4):478-94. doi: 10.1007/s002210050484.
Interactions between proprioceptive and vestibular inputs contributing to the generation of balance corrections may vary across muscles depending on the availability of sensory information at centres initiating and modulating muscle synergies, and the efficacy with which the muscle action can prevent a fall. Information which is not available from one sensory system may be obtained by switching to another. Alternatively, interactions between sensory systems and the muscle to which this interaction is targeted may be fixed during neural development and not switchable. To investigate these different concepts, balance corrections with three different sets of proprioceptive trigger signals were examined under eyes-open and eyes-closed conditions in the muscles of normal subjects and compared with those of subjects with bilateral peripheral vestibular loss. The different sets of early proprioceptive inputs were obtained by employing three combinations of support surface rotation and translation, for which ankle inputs were nulled, normal or enhanced, the knees were either locked or in flexion, and the trunk was either in flexion or extension. Three types of proprioceptive and vestibulospinal interactions were identified in muscles responses. These interactions were typified by the responses of triceps surae, quadriceps, and paraspinal muscles. The amplitudes of stretch responses at 50 ms after the onset of ankle flexion in triceps surae muscles were related to the velocity of ankle stretch. The amplitude of balance-correcting responses at 100 ms corresponded more with stretch of the biarticular gastrocnemius when the knee was re-extended at 60 ms. Absent stretch reflexes at 50 ms in triceps surae with nulled ankle inputs caused a minor, 12-ms delay in the onset of balance-correcting responses in triceps surae muscles. Vestibular loss caused no change in the amplitude of balance-correcting responses, but a negligible decrease in onset latency in triceps surae even with nulled ankle inputs. Stretch responses in quadriceps at 80 ms increased with the velocity of knee flexion but were overall lower in amplitude in vestibular loss subjects. Balance-correcting responses in quadriceps had amplitudes which were related to the directions of initial trunk movements, were still present when knee inputs were negligible and were also altered after vestibular loss. Stretch and unloading responses in paraspinals at 80 ms were consistent with the direction of initial trunk flexion and extension. Subsequent balance-correcting responses in paraspinals were delayed 20 ms in onset and altered in amplitude by vestibular loss. The changes in the amplitudes of ankle (tibialis anterior), knee (quadriceps) and trunk (paraspinal) muscle responses with vestibular loss affected the amplitudes and timing of trunk angular velocities, requiring increased stabilizing tibialis anterior, paraspinal and trapezius responses post 240 ms as these subjects attempted to remain upright. The results suggest that trunk inputs provide an ideal candidate for triggering balance corrections as these would still be present when vestibular, ankle and knee inputs are absent. The disparity between the amplitudes of stretch reflex and automatic balance-correcting responses in triceps surae and the insignificant alteration in the timing of balance-correcting responses in these muscles with nulled ankle inputs indicates that ankle inputs do not trigger balance corrections. Furthermore, modulation of balance corrections normally performed by vestibular inputs in some but not all muscles is not achieved by switching to another sensory system on vestibular loss. We postulate that a confluence of trunk and upper-leg proprioceptive input establishes the basic timing of automatic, triggered balance corrections which is then preferentially weighted by vestibular modulation in muscles that prevent falling. (ABSTRACT TRUNCATED)
本体感觉与前庭输入之间的相互作用有助于产生平衡校正,这种相互作用在不同肌肉中可能会有所不同,这取决于启动和调节肌肉协同作用的中枢的感觉信息可用性,以及肌肉动作预防跌倒的效果。一个感觉系统无法提供的信息可以通过切换到另一个系统来获取。或者,感觉系统与该相互作用所针对的肌肉之间的相互作用可能在神经发育过程中就已固定,无法切换。为了研究这些不同的概念,在睁眼和闭眼条件下,对正常受试者肌肉中三种不同的本体感觉触发信号组进行了平衡校正检查,并与双侧外周前庭丧失的受试者进行了比较。通过采用支撑面旋转和平移的三种组合来获得不同的早期本体感觉输入组,对于这些组合,踝关节输入被消除、正常或增强,膝盖要么锁定要么屈曲,躯干要么屈曲要么伸展。在肌肉反应中识别出三种类型的本体感觉与前庭脊髓相互作用。这些相互作用以腓肠肌、股四头肌和椎旁肌的反应为典型。腓肠肌在踝关节屈曲开始后50毫秒时的拉伸反应幅度与踝关节拉伸速度有关。当膝盖在60毫秒时重新伸展时,100毫秒时的平衡校正反应幅度与双关节腓肠肌的拉伸更相关。踝关节输入被消除时,腓肠肌在50毫秒时没有拉伸反射,导致腓肠肌肌肉平衡校正反应开始延迟12毫秒。前庭丧失不会导致平衡校正反应幅度的变化,但即使踝关节输入被消除,腓肠肌的起始潜伏期也会有可忽略不计的缩短。股四头肌在80毫秒时的拉伸反应随着膝盖屈曲速度的增加而增加,但在前庭丧失受试者中总体幅度较低。股四头肌的平衡校正反应幅度与初始躯干运动方向有关,当膝盖输入可忽略不计时仍然存在,并且在前庭丧失后也会改变。椎旁肌在80毫秒时的拉伸和卸载反应与初始躯干屈伸方向一致。随后椎旁肌的平衡校正反应开始延迟20毫秒,并且在前庭丧失后幅度改变。前庭丧失时,踝关节(胫骨前肌)、膝盖(股四头肌)和躯干(椎旁肌)肌肉反应幅度的变化影响了躯干角速度的幅度和时间,当这些受试者试图保持直立时,需要在240毫秒后增加胫骨前肌、椎旁肌和斜方肌的稳定反应。结果表明,躯干输入为触发平衡校正提供了一个理想的候选因素,因为当前庭、踝关节和膝盖输入不存在时,这些输入仍然会存在。腓肠肌拉伸反射幅度与自动平衡校正反应之间的差异以及踝关节输入被消除时这些肌肉平衡校正反应时间的微小变化表明,踝关节输入不会触发平衡校正。此外,在前庭丧失时,并非所有肌肉中通常由前庭输入执行的平衡校正调节都能通过切换到另一个感觉系统来实现。我们推测,躯干和大腿本体感觉输入的汇合确定了自动触发平衡校正的基本时间,然后在预防跌倒的肌肉中,前庭调节会优先对其进行加权。