Segal M, Barker J L
J Neurophysiol. 1984 Jun;51(6):1409-33. doi: 10.1152/jn.1984.51.6.1409.
Two-electrode voltage-clamp methodology was used to analyze voltage-dependent ionic conductances in 81 rat hippocampal neurons grown in culture for 4-6 wk. Pyramidal and multipolar cells with 15- to 20-micron-diameter cell bodies were impaled with two independent KCl electrodes. The cells had resting potentials of -30 to -60 mV and an average input resistance of about 30 M omega. A depolarizing command applied to a cell maintained in normal medium invariably evoked a fast (2-10 ms) inward current that saturated the current-passing capacity of the system. This was blocked in a reversible manner by application of tetrodotoxin (TTX) (0.1-1.0 microM) near the recorded cell. In the presence of TTX, a depolarizing command evoked a rapidly rising (3-5 ms), rapidly decaying (25 ms) transient outward current reminiscent of "IA" reported in molluscan neurons. This was followed by a more slowly activating (approximately 100 ms) outward current response of greater amplitude that decayed with a time constant of about 2-3 s. These properties resemble those associated with the K+ conductance, IK, underlying delayed rectification described in many excitable membranes. IK was blocked by extracellular application of tetraethylammonium (TEA) but was insensitive to 4-aminopyridine (4-AP) at concentrations that effectively eliminated IA. IA, in turn, was only marginally depressed by TEA. Unlike IK, IA was completely inactivated when the membrane was held at potentials positive to -50 mV. Inactivation was completely removed by conditioning hyperpolarization at -90 mV. A brief hyperpolarizing pulse (10 ms) was sufficient to remove 95% of the inactivation. IA activated on commands to potentials more positive than -50 mV. The inversion potential of the ionic conductance underlying IA and IK was in the range of the K+ equilibrium potential, EK, as measured by the inversion of tail currents; and this potential was shifted in a depolarizing direction by elevated [K+]0. Thus, both current species reflect activation of membrane conductance to K+ ions. Hyperpolarizing commands from resting potentials revealed a time- and voltage-dependent slowly developing inward current in the majority of cells studied. This membrane current was observed in cells exhibiting "anomalous rectification" and was therefore labeled IAR. It was activated at potentials negative to -70 mV with a time constant of 100-200 ms and was not inactivated. A return to resting potential revealed a tail current that disappeared at about EK. IAR was blocked by extracellular CS+ and was enhanced by elevating [K+]0. It thus appears to be carried by inward movement of K+ ions.(ABSTRACT TRUNCATED AT 400 WORDS)
采用双电极电压钳技术分析了81个培养4 - 6周的大鼠海马神经元的电压依赖性离子电导。用两个独立的氯化钾电极刺入胞体直径为15 - 20微米的锥体细胞和多极细胞。这些细胞的静息电位为 - 30至 - 60 mV,平均输入电阻约为30 MΩ。对处于正常培养基中的细胞施加去极化指令,总是会诱发一个快速(2 - 10毫秒)的内向电流,该电流使系统的电流通过能力饱和。在记录细胞附近施加河豚毒素(TTX)(0.1 - 1.0 microM)可使其以可逆方式阻断。在TTX存在的情况下,去极化指令诱发一个快速上升(3 - 5毫秒)、快速衰减(25毫秒)的瞬时外向电流,这让人想起软体动物神经元中报道的“IA”电流。随后是一个激活较慢(约100毫秒)、幅度更大的外向电流反应,其衰减时间常数约为2 - 3秒。这些特性类似于许多可兴奋膜中描述的与延迟整流相关的钾离子电导IK。IK可被细胞外应用四乙铵(TEA)阻断,但在有效消除IA的浓度下对4 - 氨基吡啶(4 - AP)不敏感。反过来,IA仅被TEA轻微抑制。与IK不同,当膜电位保持在高于 - 50 mV时,IA完全失活。通过在 - 90 mV进行预处理超极化可完全消除失活。一个短暂的超极化脉冲(10毫秒)足以消除95%的失活。IA在指令电位高于 - 50 mV时激活。通过尾电流反转测量,IA和IK所基于的离子电导的反转电位在钾离子平衡电位EK范围内;并且该电位因[K⁺]₀升高而向去极化方向移动。因此,这两种电流都反映了膜对钾离子电导的激活。从静息电位开始的超极化指令在大多数研究的细胞中揭示了一种时间和电压依赖性的缓慢发展的内向电流。这种膜电流在表现出“反常整流”的细胞中观察到,因此被标记为IAR。它在电位低于 - 70 mV时以100 - 200毫秒的时间常数激活,并且不会失活。回到静息电位会揭示一个在约EK处消失的尾电流。IAR被细胞外的CS⁺阻断,并因[K⁺]₀升高而增强。因此,它似乎是由钾离子的内向移动所携带。(摘要截断于400字)