Weidner C, Schmidt R, Schmelz M, Torebjork H E, Handwerker H O
Department of Physiology and Experimental Pathophysiology, University of Erlangen/Nürnberg, Germany.
J Physiol. 2003 Mar 15;547(Pt 3):931-40. doi: 10.1113/jphysiol.2002.028712. Epub 2003 Feb 7.
Recordings of single human peroneal C-fibres and rat saphenous C-fibres confirm two different patterns of conduction at branching points. In general, an action potential (AP) arising from one terminal branch may be propagated not only centrally, but also antidromically into the other branches of the terminal arborisation. If a stimulus activates several converging branches of one unit, at each branching point only the AP arriving first from the simultaneously activated daughter branches will be propagated centrally, resetting the slower branches. However, occasionally a single electrical stimulus may evoke a double response in the parent axon. In this case, these two responses apparently originate from different terminal branches and require unidirectional conduction block to prevent the faster AP from invading and resetting the slower-conducting terminal. This conclusion is supported by the notion that when such a double response occurs, both responses immediately show additional activity-dependent slowing of the conduction velocity due to frequency increase in the parent axon (two spikes per stimulus, one from each of the two excited branches). A comparable discharge pattern in the stem axon can be induced by repetitive paired stimulation of one terminal branch. Then the slowing is induced by the doubled frequency along the whole nerve fibre including the terminal branch. Since in this case not only the stem axon, but also the terminal branches carry two spikes per pulse, activity-dependent slowing is predictably more pronounced. Unidirectional block thus provides insight into the differential amount of activity-dependent slowing (and hence postexcitatory hyperpolarisation) in the stem axon and terminal branches of cutaneous C-fibres. This comparison reveals that more than two-thirds of the slowing can be attributed to the terminal branches, since it is two- to fourfold that observed during double stimulation as compared with the unidirectional block condition. This indicates that the terminal branches are equipped with membrane proteins that are different from those of the parent axon.
对单根人体腓总神经C纤维和大鼠隐神经C纤维的记录证实了分支点处存在两种不同的传导模式。一般来说,源自一个终末分支的动作电位(AP)不仅可以向中枢传播,还可以逆向传入终末分支树的其他分支。如果一个刺激激活了一个单位的几个汇聚分支,在每个分支点,只有首先从同时激活的子分支到达的AP会向中枢传播,从而重置较慢的分支。然而,偶尔单个电刺激可能会在母轴突中引发双重反应。在这种情况下,这两种反应显然源自不同的终末分支,并且需要单向传导阻滞来防止较快的AP侵入并重置传导较慢的终末分支。这一结论得到以下观点的支持:当出现这种双重反应时,由于母轴突中频率增加(每个刺激两个尖峰,分别来自两个兴奋分支中的每一个),两种反应都会立即出现额外的活动依赖性传导速度减慢。通过对一个终末分支进行重复配对刺激,可以在干轴突中诱导出类似的放电模式。然后,沿着包括终末分支在内的整个神经纤维,频率加倍会诱导出减慢。由于在这种情况下,不仅干轴突,而且终末分支每个脉冲都携带两个尖峰,因此活动依赖性减慢预计会更加明显。因此,单向阻滞为深入了解皮肤C纤维的干轴突和终末分支中活动依赖性减慢(以及因此的兴奋后超极化)的差异程度提供了线索。这种比较表明,超过三分之二的减慢可归因于终末分支,因为与单向阻滞条件相比,在双重刺激期间观察到的减慢是其两到四倍。这表明终末分支配备了与母轴突不同的膜蛋白。