D. Farina, Medical University Göttingen, Von-Siebold-Str. 4,37075 Göttingen, Germany Göttingen 37075 Germany.
J Physiol. 2013 Dec 15;591(24):6139-56. doi: 10.1113/jphysiol.2013.262477. Epub 2013 Oct 21.
It is commonly assumed that the orderly recruitment of motor units by size provides a functional advantage for the performance of movements compared with a random recruitment order. On the other hand, the excitability of a motor neuron depends on its size and this is intrinsically linked to its innervation number. A range of innervation numbers among motor neurons corresponds to a range of sizes and thus to a range of excitabilities ordered by size. Therefore, if the excitation drive is similar among motor neurons, the recruitment by size is inevitably due to the intrinsic properties of motor neurons and may not have arisen to meet functional demands. In this view, we tested the assumption that orderly recruitment is necessarily beneficial by determining if this type of recruitment produces optimal motor output. Using evolutionary algorithms and without any a priori assumptions, the parameters of neuromuscular models were optimized with respect to several criteria for motor performance. Interestingly, the optimized model parameters matched well known neuromuscular properties, but none of the optimization criteria determined a consistent recruitment order by size unless this was imposed by an association between motor neuron size and excitability. Further, when the association between size and excitability was imposed, the resultant model of recruitment did not improve the motor performance with respect to the absence of orderly recruitment. A consistent observation was that optimal solutions for a variety of criteria of motor performance always required a broad range of innervation numbers in the population of motor neurons, skewed towards the small values. These results indicate that orderly recruitment of motor units in itself does not provide substantial functional advantages for motor control. Rather, the reason for its near-universal presence in human movements is that motor functions are optimized by a broad range of innervation numbers.
人们普遍认为,与随机募集顺序相比,按大小顺序募集运动单位为运动表现提供了一种功能优势。另一方面,运动神经元的兴奋性取决于其大小,而这与它的支配数量内在相关。运动神经元的支配数量范围对应于一系列大小,从而对应于按大小排序的兴奋性范围。因此,如果运动神经元之间的兴奋驱动力相似,那么按大小募集必然是由于运动神经元的内在特性所致,而不是为了满足功能需求而产生的。在这种观点下,我们通过确定这种募集方式是否能产生最佳的运动输出,来检验募集顺序必然有益的假设。使用进化算法,并且没有任何先验假设,我们针对运动表现的几个标准,对神经肌肉模型的参数进行了优化。有趣的是,优化后的模型参数与众所周知的神经肌肉特性非常吻合,但没有任何优化标准能确定一致的按大小募集顺序,除非这种顺序是由运动神经元大小和兴奋性之间的关联决定的。此外,当大小和兴奋性之间的关联被强加时,按大小募集的结果模型并不能改善运动表现,而无需按顺序募集。一个一致的观察结果是,各种运动表现标准的最优解总是需要运动神经元群体中支配数量的广泛范围,偏向于小值。这些结果表明,运动单位的按顺序募集本身并不能为运动控制提供实质性的功能优势。相反,它在人类运动中几乎普遍存在的原因是,通过广泛的支配数量,运动功能得到了优化。