Fuglevand A J, Macefield V G, Bigland-Ritchie B
John B. Pierce Laboratory, New Haven, Connecticut 06519, USA.
J Neurophysiol. 1999 Apr;81(4):1718-29. doi: 10.1152/jn.1999.81.4.1718.
Modulation of motor unit activation rate is a fundamental process by which the mammalian nervous system encodes muscle force. To identify how rate coding of force may change as a consequence of fatigue, intraneural microstimulation of motor axons was used to elicit twitch and force-frequency responses before and after 2 min of intermittent stimulation (40-Hz train for 330 ms, 1 train/s) in single motor units of human long finger flexor muscles and intrinsic hand muscles. Before fatigue, two groups of units could be distinguished based on the stimulus frequency needed to elicit half-maximal force; group 1 (n = 8) required 9.1 +/- 0.5 Hz (means +/- SD), and group 2 (n = 5) required 15.5 +/- 1.1 Hz. Twitch contraction times were significantly different between these two groups (group 1 = 66. 5 ms; group 2 = 45.9 ms). Overall 18% of the units were fatigue resistant [fatigue index (FI) > 0.75], 64% had intermediate fatigue sensitivity (0.25 </= FI </= 0.75), and 18% were fatigable (FI < 0. 25). However, fatigability and tetanic force were not significantly different among groups. Therefore unlike findings in some other mammals, fast-contracting motor units were neither stronger nor more susceptible to fatigue than slowly contracting units. Fatigue, however, was found to be greatest in those units that initially exerted the largest forces. Despite significant slowing of contractile responses, fatigue caused the force-frequency relation to become displaced toward higher frequencies (44 +/- 41% increase in frequency for half-maximal force). Moreover, the greatest shift in the force-frequency relation occurred among those units exhibiting the largest force loss. A selective deficit in force at low frequencies of stimulation persisted for several minutes after the fatigue task. Overall, these findings suggest that with fatigue higher activation rates must be delivered to motor units to maintain the same relative level of force. Questions regarding classification of motor units and possible mechanisms by which fatigue-related slowing might coexist with a shift in the force-frequency curve toward higher frequencies are discussed.
运动单位激活率的调节是哺乳动物神经系统编码肌肉力量的一个基本过程。为了确定疲劳后力量的频率编码如何变化,在人长指屈肌和手部固有肌的单个运动单位中,在间歇性刺激(40赫兹串刺激330毫秒,每秒1串)2分钟前后,通过对运动轴突进行神经内微刺激来引发抽搐和力量-频率反应。在疲劳前,根据引发半最大力量所需的刺激频率可区分出两组运动单位;第1组(n = 8)需要9.1±0.5赫兹(平均值±标准差),第2组(n = 5)需要15.5±1.1赫兹。这两组的抽搐收缩时间有显著差异(第1组 = 66.5毫秒;第2组 = 45.9毫秒)。总体而言,18%的运动单位抗疲劳[疲劳指数(FI)> 0.75],64%具有中等疲劳敏感性(0.25≤FI≤0.75),18%易疲劳(FI < 0.25)。然而,各组之间的易疲劳性和强直力量并无显著差异。因此,与其他一些哺乳动物的研究结果不同,快速收缩的运动单位并不比缓慢收缩的单位更强壮,也不比其更易疲劳。然而,发现最初施加最大力量的那些运动单位疲劳程度最大。尽管收缩反应明显减慢,但疲劳导致力量-频率关系向更高频率偏移(半最大力量时频率增加44±41%)。此外,力量-频率关系的最大偏移发生在力量损失最大的那些运动单位中。在疲劳任务后,低频刺激时的力量选择性缺陷持续了几分钟。总体而言,这些发现表明,随着疲劳的出现,必须向运动单位传递更高的激活率以维持相同的相对力量水平。文中还讨论了关于运动单位分类以及疲劳相关的收缩减慢可能与力量-频率曲线向更高频率偏移同时存在的可能机制的问题。