Ting L H, Kautz S A, Brown D A, Zajac F E
Rehabilitation Research and Development Center (153), Veterans Affairs Palo Alto Health Care System, Palo Alto, California 94304, USA.
J Neurophysiol. 1999 Feb;81(2):544-51. doi: 10.1152/jn.1999.81.2.544.
Computer simulations of pedaling have shown that a wide range of pedaling tasks can be performed if each limb has the capability of executing six biomechanical functions, which are arranged into three pairs of alternating antagonistic functions. An Ext/Flex pair accelerates the limb into extension or flexion, a Plant/Dorsi pair accelerates the foot into plantarflexion or dorsiflexion, and an Ant/Post pair accelerates the foot anteriorly or posteriorly relative to the pelvis. Because each biomechanical function (i.e., Ext, Flex, Plant, Dorsi, Ant, or Post) contributes to crank propulsion during a specific region in the cycle, phasing of a muscle is hypothesized to be a consequence of its ability to contribute to one or more of the biomechanical functions. Analysis of electromyogram (EMG) patterns has shown that this biomechanical framework assists in the interpretation of muscle activity in healthy and hemiparetic subjects during forward pedaling. Simulations show that backward pedaling can be produced with a phase shift of 180 degrees in the Ant/Post pair. No phase shifts in the Ext/Flex and Plant/Dorsi pairs are then necessary. To further test whether this simple yet biomechanically viable strategy may be used by the nervous system, EMGs from 7 muscles in 16 subjects were measured during backward as well as forward pedaling. As predicted, phasing in vastus medialis (VM), tibialis anterior (TA), medial gastrocnemius (MG), and soleus (SL) were unaffected by pedaling direction, with VM and SL contributing to Ext, MG to Plant, and TA to Dorsi. In contrast, phasing in biceps femoris (BF) and semimembranosus (SM) were affected by pedaling direction, as predicted, compatible with their contribution to the directionally sensitive Post function. Phasing of rectus femoris (RF) was also affected by pedaling direction; however, its ability to contribute to the directionally sensitive Ant function may only be expressed in forward pedaling. RF also contributed significantly to the directionally insensitive Ext function in both forward and backward pedaling. Other muscles also appear to have contributed to more than one function, which was especially evident in backward pedaling (i.e. , BF, SM, MG, and TA to Flex). We conclude that the phasing of only the Ant and Post biomechanical functions are directionally sensitive. Further, we suggest that task-dependent modulation of the expression of the functions in the motor output provides this biomechanics-based neural control scheme with the capability to execute a variety of lower limb tasks, including walking.
对蹬踏动作的计算机模拟表明,如果每个肢体都具备执行六种生物力学功能的能力,那么就能完成范围广泛的蹬踏任务,这六种功能可分为三对交替的拮抗功能。伸展/屈曲对使肢体加速伸展或屈曲,跖屈/背屈对使足部加速跖屈或背屈,前/后对使足部相对于骨盆向前或向后加速。由于每个生物力学功能(即伸展、屈曲、跖屈、背屈、前、或后)在周期中的特定阶段对曲柄推进有贡献,因此推测肌肉的相位是其对一种或多种生物力学功能有贡献能力的结果。肌电图(EMG)模式分析表明,这种生物力学框架有助于解释健康受试者和偏瘫受试者在向前蹬踏过程中的肌肉活动。模拟显示,通过前/后对中180度的相位偏移可以产生向后蹬踏。此时伸展/屈曲对和跖屈/背屈对无需相位偏移。为了进一步测试这种简单但在生物力学上可行的策略是否可能被神经系统使用,在16名受试者向后和向前蹬踏过程中测量了7块肌肉的肌电图。正如预测的那样,股内侧肌(VM)、胫骨前肌(TA)、腓肠肌内侧头(MG)和比目鱼肌(SL)的相位不受蹬踏方向影响,VM和SL对伸展有贡献,MG对跖屈有贡献,TA对背屈有贡献。相比之下,正如预测的那样,股二头肌(BF)和半膜肌(SM)的相位受蹬踏方向影响,这与其对方向敏感的后功能的贡献相符。股直肌(RF)的相位也受蹬踏方向影响;然而,其对方向敏感的前功能的贡献能力可能仅在向前蹬踏时表现出来。RF在向前和向后蹬踏中对方向不敏感的伸展功能也有显著贡献。其他肌肉似乎也对不止一种功能有贡献,这在向后蹬踏时尤为明显(即BF、SM、MG和TA对屈曲)。我们得出结论,只有前和后生物力学功能的相位对方向敏感。此外,我们认为运动输出中功能表达的任务依赖性调制为这种基于生物力学的神经控制方案提供了执行各种下肢任务(包括行走)的能力。