神经传导热的热力学框架
The Heat of Nervous Conduction: A Thermodynamic Framework
摘要：
神经传导的早期记录揭示了与电信号相关的显著的热特征。所观察到的热量产生和随后的吸收是由动作电位传导期间在细胞膜水平发生的物理化学过程引起的。特别是，可逆释放的电能储存为跨细胞膜的电位差，似乎是一个简单而一致的解释热生产，提出了“冷凝器理论。“然而，冷凝器理论还没有被分析超出细胞膜和平行板电容器之间的类比，即冷凝器，它不能解释热信号的大小。在这项工作中，我们使用一个详细的静电模型的细胞膜重温电容器理论。我们推导出的自由能和熵的变化与膜的去极化的动作电位，这给出了一个直接的措施，由神经元产生和吸收的热量的表达式。我们展示了膜两侧的表面电荷密度如何影响能量变化。最后，考虑到一个典型的动作电位，我们表明，如果膜持有的表面电荷的偏置，使膜的内侧是0.05 C m $^{-2}$更负比外侧，该模型预测的热量的大小达到实验值的范围。基于我们的研究，我们确定了在神经传导过程中，神经元产生和吸收热量的主要机制是膜的电能变化。
英文摘要：
Early recordings of nervous conduction revealed a notable thermal signature associated with the electrical signal. The observed production and subsequent absorption of heat arise from physicochemical processes that occur at the cell membrane level during the conduction of the action potential. In particular, the reversible release of electrical energy stored as a difference of potential across the cell membrane appears as a simple yet consistent explanation for the heat production, as proposed in the "Condenser Theory." However, the Condenser Theory has not been analyzed beyond the analogy between the cell membrane and a parallel-plate capacitor, i.e. a condenser, which cannot account for the magnitude of the heat signature. In this work, we use a detailed electrostatic model of the cell membrane to revisit the Condenser Theory. We derive expressions for free energy and entropy changes associated with the depolarization of the membrane by the action potential, which give a direct measure of the heat produced and absorbed by neurons. We show how the density of surface charges on both sides of the membrane impacts the energy changes. Finally, considering a typical action potential, we show that if the membrane holds a bias of surface charges, such that the internal side of the membrane is 0.05 C m$^{-2}$ more negative than the external side, the size of the heat predicted by the model reaches the range of experimental values. Based on our study, we identify the change in electrical energy of the membrane as the primary mechanism of heat production and absorption by neurons during nervous conduction.
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