London M, Meunier C, Segev I
Department of Neurobiology, Institute of Life Sciences and Center for Neural Computation, Hebrew University, Jerusalem 91904, Israel.
J Neurosci. 1999 Oct 1;19(19):8219-33. doi: 10.1523/JNEUROSCI.19-19-08219.1999.
In recent years it became clear that dendrites possess a host of ion channels that may be distributed nonuniformly over their membrane surface. In cortical pyramids, for example, it was demonstrated that the resting membrane conductance G(m)(x) is higher (the membrane is "leakier") at distal dendritic regions than at more proximal sites. How does this spatial nonuniformity in G(m)(x) affect the input-output function of the neuron? The present study aims at providing basic insights into this question. To this end, we have analytically studied the fundamental effects of membrane non-uniformity in passive cable structures. Keeping the total membrane conductance over a given modeled structure fixed (i.e., a constant number of passive ion channels), the classical case of cables with uniform membrane conductance is contrasted with various nonuniform cases with the following general conclusions. (1) For cylindrical cables with "sealed ends," monotonic increase in G(m)(x) improves voltage transfer from the input location to the soma. The steeper the G(m)(x), the larger the improvement. (2) This effect is further enhanced when the stimulation is distal and consists of a synaptic input rather than a current source. (3) Any nonuniformity in G(m)(x) decreases the electrotonic length, L, of the cylinder. (4) The system time constant tau(0) is larger in the nonuniform case than in the corresponding uniform case. (5) When voltage transients relax with tau(0), the dendritic tree is not isopotential in the nonuniform case, at variance with the uniform case. The effect of membrane nonuniformity on signal transfer in reconstructed dendritic trees and on the I/f relation of the neuron is also considered, and experimental methods for assessing membrane nonuniformity in dendrites are discussed.
近年来,越来越清楚的是,树突拥有大量离子通道,这些通道可能在其膜表面非均匀分布。例如,在皮质锥体神经元中,已证明静息膜电导G(m)(x)在树突远端区域比在更近端部位更高(膜更“易漏”)。G(m)(x)的这种空间非均匀性如何影响神经元的输入-输出功能?本研究旨在为这个问题提供基本见解。为此,我们对被动电缆结构中膜非均匀性的基本影响进行了分析研究。在给定建模结构上保持总膜电导固定(即被动离子通道数量恒定),将具有均匀膜电导的电缆的经典情况与各种非均匀情况进行对比,得出以下一般结论。(1) 对于“封闭端”的圆柱形电缆,G(m)(x)的单调增加改善了从输入位置到胞体的电压传递。G(m)(x)越陡峭,改善越大。(2) 当刺激是远端的且由突触输入而非电流源组成时,这种效应会进一步增强。(3) G(m)(x)的任何非均匀性都会减小圆柱体的电紧张长度L。(4) 非均匀情况下的系统时间常数tau(0)比相应均匀情况下的更大。(5) 当电压瞬变以tau(0)松弛时,在非均匀情况下树突树不是等电位的,这与均匀情况不同。还考虑了膜非均匀性对重建树突树中信号传递和神经元I/f关系的影响,并讨论了评估树突中膜非均匀性的实验方法。