Laboratory of Integrative Physiology, Faculty of Integrated Arts and Sciences, Hiroshima University, Higashi-Hiroshima, 739-8521, Japan,
J Comp Physiol B. 2013 Dec;183(8):1015-21. doi: 10.1007/s00360-013-0770-3. Epub 2013 Jun 8.
By developing a new in vivo method to evaluate the esophageal closure, which reflects inhibition of swallowing, we demonstrate that the vagal X1 branch projected from the glossopharyngeal-vagal motor complex (GVC) controls the upper esophageal sphincter (UES) muscle directly. Although eel vagal nerve consisted of five branches, other branches (X2, X3, X4 and X5) did not influence the esophageal pressure. When the X1 nerve branch was stimulated electrically, the balloon pressure in the UES area increased with optimum frequency of 20 Hz. Since similar optimum frequency was observed both in the pithed eel and in the isolated UES preparation, such characteristic of X1 nerve is not due to anesthetic used during experiment. As the isolated UES preparation consists of muscle cells and nerve terminals, and as the optimum frequency of the nerve terminal is identical with that of the X1 branch, it is most likely that the X1 nerve branch is identical with the nerve terminals within the UES preparation. On the other hand, since the GVC neurons fire spontaneously at around 20 Hz, the optimum frequency of 20 Hz means that the eel UES is usually closed vigorously and relaxed only when the GVC neuron is inactivated. The effect of X1 stimulation was inhibited by curare, but not by atropine, indicating that the X1 nerve branch releases acetylcholine, which acts on the nicotinic receptor on the UES striated muscle. Beside vagal nerve X1 branch, spinal nerve SN2, SN3 and SN4 also contributed to the UES closure, but SN1 did not influence the UES movement. However, since the efficacy of these spinal nerve stimulations is about 1/10 of that by vagal X1 branch, the eel UES may be controlled primarily by a vagal nerve X1 branch, and secondarily by spinal nerves (SN2, SN3 and SN4).
通过开发一种新的体内方法来评估食管闭合,反映吞咽抑制,我们证明了从舌咽-迷走运动复合体(GVC)投射的迷走神经 X1 分支直接控制食管上括约肌(UES)肌肉。尽管鳗鱼的迷走神经由五个分支组成,但其他分支(X2、X3、X4 和 X5)不会影响食管压力。当 X1 神经分支受到电刺激时,UES 区域的气球压力会随着 20 Hz 的最佳频率增加。由于在去脑鳗鱼和分离的 UES 制剂中都观察到了类似的最佳频率,因此 X1 神经的这种特征不是由于实验中使用的麻醉剂所致。由于分离的 UES 制剂由肌肉细胞和神经末梢组成,并且神经末梢的最佳频率与 X1 分支相同,因此 X1 神经分支很可能与 UES 制剂内的神经末梢相同。另一方面,由于 GVC 神经元以大约 20 Hz 的频率自发放电,因此 20 Hz 的最佳频率意味着鳗鱼 UES 通常会剧烈关闭,只有当 GVC 神经元失活时才会放松。X1 刺激的效果被箭毒抑制,但不受阿托品抑制,表明 X1 神经分支释放乙酰胆碱,乙酰胆碱作用于 UES 横纹肌上的烟碱受体。除了迷走神经 X1 分支外,脊神经 SN2、SN3 和 SN4 也有助于 UES 关闭,但 SN1 不会影响 UES 运动。然而,由于这些脊神经刺激的效果约为 X1 分支的 1/10,因此鳗鱼 UES 可能主要由迷走神经 X1 分支控制,其次是由脊神经(SN2、SN3 和 SN4)控制。