非线性对共振神经元非对称电压响应的影响
Asymmetrical voltage response in resonant neurons shaped by nonlinearities
摘要：
神经元的常规阻抗分布可以识别谐振的存在和神经元响应对振荡输入的其他特性，例如非线性响应放大，但是它不能区分其他非线性特性，例如电压响应包络形状的不对称性。实验观察表明，神经元对振荡输入的响应在不同的频带中优先增强电压包络的上部或下部。这些不对称的电压响应出现在神经元模型中，当它被提交到足够高的振幅振荡电流的可变频率。我们展示了如何与不同的离子电流或存在于模型中的非线性捕获其电压方程导致不对称的响应，以及如何高振幅振荡电流强调这种响应。我们提出了一个几何解释的现象，其中不对称的结果不仅从非线性的激活曲线，但也从非线性捕获的零倾在相平面图和系统的时间尺度分离。此外，我们确定了一个意想不到的频率依赖性模式，它在这些电流的门控变量中发展，并且是系统中强非线性的产物，正如我们通过操纵激活曲线参数来控制这种行为所示。本文报道的结果揭示了脑内神经元处理振荡信息的离子机制。
英文摘要：
The conventional impedance profile of a neuron can identify the presence of resonance and other properties of the neuronal response to oscillatory inputs, such as nonlinear response amplifications, but it cannot distinguish other nonlinear properties such as asymmetries in the shape of the voltage response envelope. Experimental observations have shown that the response of neurons to oscillatory inputs preferentially enhances either the upper or lower part of the voltage envelope in different frequency bands. These asymmetric voltage responses arise in a neuron model when it is submitted to high enough amplitude oscillatory currents of variable frequencies. We show how the nonlinearities associated to different ionic currents or present in the model as captured by its voltage equation lead to asymmetrical response and how high amplitude oscillatory currents emphasize this response. We propose a geometrical explanation for the phenomenon where asymmetries result not only from nonlinearities in their activation curves but also from nonlinearites captured by the nullclines in the phase-plane diagram and from the system's time-scale separation. In addition, we identify an unexpected frequency-dependent pattern which develops in the gating variables of these currents and is a product of strong nonlinearities in the system as we show by controlling such behavior by manipulating the activation curve parameters. The results reported in this paper shed light on the ionic mechanisms by which brain embedded neurons process oscillatory information.
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