Ekeberg O, Grillner S
Department of Numerical Analysis and Computing Science, Royal Institute of Technology, Stockholm, Sweden.
Philos Trans R Soc Lond B Biol Sci. 1999 May 29;354(1385):895-902. doi: 10.1098/rstb.1999.0441.
The neuronal generation of vertebrate locomotion has been extensively studied in the lamprey. Models at different levels of abstraction are being used to describe this system, from abstract nonlinear oscillators to interconnected model neurons comprising multiple compartments and a Hodgkin-Huxley representation of the most relevant ion channels. To study the role of sensory feedback by simulation, it eventually also becomes necessary to incorporate the mechanical movements in the models. By using simplifying models of muscle activation, body mechanics, counteracting water forces, and sensory feedback through stretch receptors and vestibular organs, we have been able to close the feedback loop to enable studies of the interaction between the neuronal and the mechanical systems. The neuromechanical simulations reveal that the currently known network is sufficient for generating a whole repertoire of swimming patterns. Swimming at different speeds and with different wavelengths, together with the performance of lateral turns can all be achieved by simply varying the brainstem input. The neuronal mechanisms behind pitch and roll manoeuvres are less clear. We have put forward a 'crossed-oscillators' hypothesis where partly separate dorsal and ventral circuits are postulated. Neuromechanical simulations of this system show that it is also capable of generating realistic pitch turns and rolls, and that vestibular signals can stabilize the posture during swimming.
脊椎动物运动的神经元生成过程已在七鳃鳗中得到广泛研究。人们使用不同抽象层次的模型来描述这个系统,从抽象的非线性振荡器到由多个隔室组成的相互连接的模型神经元,以及对最相关离子通道的霍奇金 - 赫胥黎表示。为了通过模拟研究感觉反馈的作用,最终还需要在模型中纳入机械运动。通过使用肌肉激活、身体力学、抵消水动力以及通过拉伸感受器和前庭器官进行感觉反馈的简化模型,我们得以闭合反馈回路,从而能够研究神经元系统与机械系统之间的相互作用。神经力学模拟表明,目前已知的网络足以生成完整的游泳模式库。以不同速度和不同波长游泳,以及进行侧向转弯,都可以通过简单改变脑干输入来实现。俯仰和翻滚动作背后的神经元机制尚不清楚。我们提出了一个“交叉振荡器”假说,假定存在部分分离的背侧和腹侧回路。对该系统的神经力学模拟表明,它也能够产生逼真的俯仰转弯和翻滚,并且前庭信号可以在游泳过程中稳定姿势。