Schwarz C, Möck M, Thier P
Sektion für Visuelle Sensomotorik, Neurologische Universitätsklinik Tübingen, 72076 Tubingen, Germany.
J Neurophysiol. 1997 Dec;78(6):3323-37. doi: 10.1152/jn.1997.78.6.3323.
We used a new slice preparation of rat brain stem to establish the basic membrane properties of neurons in the pontine nuclei (PN). Using standard intracellular recordings, we found that pontine cells displayed a resting membrane potential of -63 +/- 6 mV (mean +/- SD), an input resistance of 53 +/- 21 MOmega, a membrane time constant of 5.3 +/- 2.4 ms and were not spontaneously active. The current-voltage relationship of most of the PN neurons showed the characteristics of inward rectification in both depolarizing and hyperpolarizing directions. A prominent feature of the firing of pontine neurons was a marked firing rate adaptation, which eventually caused the cells to cease firing. Several types of membrane conductances possibly contribute to this feature. For one, a medium and a slow type of afterhyperpolarization (AHP) control the pattern of firing. The medium AHP was partly susceptible to blockade of calcium influx, whereas it was abolished completely by blockade of potassium channels with tetraethylammonium, indicating that it is based on at least two conductances: a calcium-dependent and a calcium-independent one. The slow AHP was carried by potassium ions and could be blocked effectively by preventing calcium influx into the cell. It was present after single spikes but was strongest after a high-frequency spike train. Calcium entry into the cell was mediated by high-threshold calcium channels that were detected by the generation of calcium spikes under blockade of potassium channels. Furthermore, the early phase of the firing rate adaptation was shown to be related to the time course of a slow, tetrodotoxin (TTX)-sensitive, persistent sodium potential, which was activated already in the subthreshold range of membrane potentials. This potential was time dependent and imposed as a depolarizing "hump" with a maximum occurring in most cases between 50 and 100 ms after stimulus onset. In the suprathreshold range, it generated plateau potentials following fast spikes, if potassium channels were blocked. After the complete adaptation of the firing rate, PN neurons were observed to display irregular fluctuations of the membrane potential, which sometimes reached firing threshold thereby eliciting an irregular low-frequency spike train. As these fluctuations could be blocked with TTX, they probably are based on the persistent sodium currents. The opposing drive in hyperpolarizing direction may be provided by strong outward currents that generated a marked outward rectification in the current-voltage relationship under TTX. In conclusion, PN neurons show complex membrane properties that are reminiscent in many ways to cerebrocortical "regular firing" neurons.
我们使用大鼠脑干的一种新切片制备方法来确定脑桥核(PN)中神经元的基本膜特性。通过标准的细胞内记录,我们发现脑桥细胞的静息膜电位为-63±6 mV(平均值±标准差),输入电阻为53±21 MΩ,膜时间常数为5.3±2.4 ms,且无自发活动。大多数脑桥核神经元的电流-电压关系在去极化和超极化方向均表现出内向整流特性。脑桥神经元放电的一个显著特征是明显的放电频率适应性,最终导致细胞停止放电。几种类型的膜电导可能促成了这一特征。其一,一种中等和一种缓慢的超极化后电位(AHP)控制着放电模式。中等AHP部分易受钙内流阻断的影响,而用四乙铵阻断钾通道可使其完全消除,这表明它至少基于两种电导:一种钙依赖性电导和一种钙非依赖性电导。缓慢AHP由钾离子介导,通过阻止钙流入细胞可有效阻断。它在单个动作电位后出现,但在高频动作电位串后最强。钙进入细胞是由高阈值钙通道介导的,在钾通道阻断情况下通过产生钙尖峰得以检测。此外,放电频率适应性的早期阶段显示与一种缓慢的、对河豚毒素(TTX)敏感的持续性钠电位的时间进程有关,该电位在膜电位的阈下范围内就已被激活。这种电位具有时间依赖性,表现为一个去极化“驼峰”,在大多数情况下,其最大值出现在刺激开始后50至100 ms之间。在阈上范围内,如果钾通道被阻断,它会在快速动作电位后产生平台电位。在放电频率完全适应后,观察到脑桥核神经元表现出膜电位的不规则波动,有时会达到放电阈值,从而引发不规则的低频动作电位串。由于这些波动可用TTX阻断,它们可能基于持续性钠电流。在超极化方向的相反驱动力可能由强大的外向电流提供,该电流在TTX存在下的电流-电压关系中产生明显的外向整流。总之,脑桥核神经元表现出复杂的膜特性,在许多方面类似于大脑皮质的“规则放电”神经元。