Department of Pharmacology-Physiology, Faculty of Medicine and Health Sciences, Université de Sherbrooke, Sherbrooke, Quebec J1H 5N4, Canada, and.
Department of Neurobiology and Anatomy, Drexel University College of Medicine, Philadelphia, Pennsylvania 19129.
J Neurosci. 2018 Apr 25;38(17):4104-4122. doi: 10.1523/JNEUROSCI.3288-17.2018. Epub 2018 Mar 21.
When the foot contacts an obstacle during locomotion, cutaneous inputs activate spinal circuits to ensure dynamic balance and forward progression. In quadrupeds, this requires coordinated reflex responses between the four limbs. Here, we investigated the patterns and phasic modulation of cutaneous reflexes in forelimb and hindlimb muscles evoked by inputs from all four limbs. Five female cats were implanted to record muscle activity and to stimulate the superficial peroneal and superficial radial nerves during locomotion. Stimulating these nerves evoked short-, mid-, and longer-latency excitatory and/or inhibitory responses in all four limbs that were phase-dependent. The largest responses were generally observed during the peak activity of the muscle. Cutaneous reflexes during mid-swing were consistent with flexion of the homonymous limb and accompanied by modification of the stance phases of the other three limbs, by coactivating flexors and extensors and/or by delaying push-off. Cutaneous reflexes during mid-stance were consistent with stabilizing the homonymous limb by delaying and then facilitating its push-off and modifying the support phases of the homolateral and diagonal limbs, characterized by coactivating flexors and extensors, reinforcing extensor activity and/or delaying push-off. The shortest latencies of homolateral and diagonal responses were consistent with fast-conducting disynaptic or trisynaptic pathways. Descending homolateral and diagonal pathways from the forelimbs to the hindlimbs had a higher probability of eliciting responses compared with ascending pathways from the hindlimbs to the forelimbs. Thus, in quadrupeds, intralimb and interlimb reflexes activated by cutaneous inputs ensure dynamic coordination of the four limbs, producing a whole-body response. The skin contains receptors that, when activated, send inputs to spinal circuits, signaling a perturbation. Rapid responses, or reflexes, in muscles of the contacted limb and opposite homologous limb help maintain balance and forward progression. Here, we investigated reflexes during quadrupedal locomotion in the cat by electrically stimulating cutaneous nerves in each of the four limbs. Functionally, responses appear to modify the trajectory or stabilize the movement of the stimulated limb while modifying the support phase of the other limbs. Reflexes between limbs are mediated by fast-conducting pathways that involve excitatory and inhibitory circuits controlling each limb. The comparatively stronger descending pathways from cervical to lumbar circuits controlling the forelimbs and hindlimbs, respectively, could serve a protective function.
当动物在运动过程中足部接触障碍物时,皮肤传入信息激活脊髓回路以确保动态平衡和前进运动。在四足动物中,这需要四肢之间协调的反射反应。在这里,我们研究了来自四肢的所有输入诱发的前肢和后肢肌肉中的皮肤反射的模式和时相调制。五只雌性猫被植入以在运动过程中记录肌肉活动并刺激腓浅神经和桡浅神经。刺激这些神经在所有四肢中诱发短、中、长潜伏期的兴奋性和/或抑制性反应,这些反应具有时相依赖性。最大的反应通常发生在肌肉活动峰值期间。在中间摆动期间的皮肤反射与同源肢体的屈曲一致,并伴随着其他三个肢体的支撑阶段的改变,通过共同激活屈肌和伸肌和/或延迟推离。在中间支撑期间的皮肤反射与稳定同源肢体一致,通过延迟然后促进其推离并改变同侧和对角肢体的支撑阶段,其特征在于共同激活屈肌和伸肌,增强伸肌活动和/或延迟推离。同侧和对角反应的最短潜伏期与快速传导的双突触或三突触通路一致。来自前肢到后肢的下行同侧和对角通路比来自后肢到前肢的上行通路更有可能引发反应。因此,在四足动物中,皮肤传入信息激活的肢体内和肢体间反射确保了四肢的动态协调,产生了全身反应。皮肤包含受体,当被激活时,会向脊髓回路发送输入,发出扰动信号。被接触肢体和相对同源肢体的肌肉中的快速反应或反射有助于维持平衡和前进运动。在这里,我们通过电刺激四肢的每个皮肤神经来研究猫的四足运动中的反射。从功能上讲,反应似乎改变了受刺激肢体的轨迹或稳定了其运动,同时改变了其他肢体的支撑阶段。肢体之间的反射由快速传导的通路介导,涉及控制每个肢体的兴奋性和抑制性回路。来自颈椎到腰椎回路的相对较强的下行通路分别控制前肢和后肢,可以起到保护作用。