Boorman G, Windhorst U, Kirmayer D
University of Calgary, Faculty of Medicine, Department of Clinical Neurosciences and Medical Physiology, Alberta, Canada.
Neuroscience. 1994 Dec;63(3):747-56. doi: 10.1016/0306-4522(94)90519-3.
A considerable number of theoretical and experimental studies have been undertaken to establish quantitative relationships between the time course of postsynaptic potentials in a neuron and the change in firing probability thereby induced. Depending on background synaptic noise level, the time course of the postsynaptic potential per se as well as its time derivative are both of importance in varying proportion. We have recently begun to study recurrent inhibitory potentials in cat hindlimb motoneurons during rhythmically varying rates of stimulation of motor axons. The amplitude-rate relationship exhibits hysteresis in that amplitudes are usually larger during augmenting than decrementing rates in the cycle. We here report results on the other important variable, that is the slope of recurrent inhibitory potential development, which need not a priori be correlated with amplitude. We found that the slope has a relation to stimulus rate similar to amplitude, so that both parameters are correlated. In pentobarbitone anaesthetized or decerebrate cats, intracellular recordings were obtained from hindlimb skeleto-motoneurons. Various hindlimb muscle nerves were prepared for electrical stimulation to elicit recurrent inhibitory potentials, with dorsal roots cut. Test stimulus patterns consisted of repetitive pulse trains whose rates varied, at modulation frequencies between 0.1 and 1.0 Hz, in one of two waveforms: triangular or sinusoidal. Modulation depths were either "full", with rates varying between a minimum of less than 10 and a maximum of around 50 pulses per s. Or they were about "half" this depth, with mean rates shifted into a "low", "medium" or "high" rate region. Recurrent inhibitory potentials were averaged with respect to stimuli occurring during different phases of the stimulation cycle. Most often when, throughout the cycle, the amplitude changed in a consistent way, so did the slopes of the inhibitory potentials. That is, when the amplitudes rhythmically declined with increasing and recovered with decreasing stimulus rate, the rate of hyperpolarization followed the same pattern. With prominent hysteresis in amplitude, a corresponding hysteresis appeared in slopes. Hence, amplitude and slopes were correlated, occasionally showing a hysteresis among themselves. To a certain extent, these results can be explained by Renshaw cell behaviour, the contribution of the Renshaw cell-motoneuron synapse being unknown and difficult to assess experimentally. For the inhibitory effect of Renshaw cells on motoneurons (and reciprocal Ia inhibitory interneurons), both its magnitude and its time course probably play an important role in determining the efficacy of counteracting local excitatory inputs. The change in slope of inhibitory potentials, and likely its underlying conductance, during cyclic motoneuron activation can be presumed to significantly contribute to the temporal pattern of discharge of motoneurons, in particular in relation to the prevention of synchronization leading to enhanced tremor.
为了建立神经元中突触后电位的时间进程与由此诱发的放电概率变化之间的定量关系,已经进行了大量的理论和实验研究。根据背景突触噪声水平,突触后电位本身的时间进程及其时间导数在不同比例下都很重要。我们最近开始研究在有节奏地改变运动轴突刺激频率时猫后肢运动神经元中的回返抑制电位。幅度 - 频率关系呈现滞后现象,即周期中增强时的幅度通常大于减弱时的幅度。我们在此报告关于另一个重要变量的结果,即回返抑制电位发展的斜率,它不必先验地与幅度相关。我们发现斜率与刺激频率的关系类似于幅度,因此这两个参数是相关的。在戊巴比妥麻醉或去大脑的猫中,从后肢骨骼肌运动神经元获得细胞内记录。准备各种后肢肌肉神经进行电刺激以诱发回返抑制电位,切断背根。测试刺激模式由重复脉冲序列组成,其频率在0.1至1.0Hz的调制频率下以两种波形之一变化:三角形或正弦形。调制深度要么是“全”,频率在每秒最少小于10次到最多约50次脉冲之间变化。要么约为这个深度的“一半”,平均频率转移到“低”、“中”或“高”频率区域。回返抑制电位相对于刺激周期不同阶段出现的刺激进行平均。最常见的是,在整个周期中,当幅度以一致的方式变化时,抑制电位的斜率也会如此。也就是说,当幅度随着刺激频率增加而有节奏地下降并随着刺激频率降低而恢复时,超极化速率遵循相同的模式。由于幅度存在明显的滞后现象,斜率中也出现了相应的滞后现象。因此,幅度和斜率是相关的,偶尔它们之间也会出现滞后现象。在一定程度上,这些结果可以用闰绍细胞的行为来解释,闰绍细胞 - 运动神经元突触的贡献尚不清楚且难以通过实验评估。对于闰绍细胞对运动神经元(以及相互的Ia抑制性中间神经元)的抑制作用,其大小和时间进程可能在决定抵消局部兴奋性输入的功效中起重要作用。在周期性运动神经元激活期间抑制电位斜率的变化,以及可能其潜在的电导变化,可以推测对运动神经元的放电时间模式有显著贡献,特别是与防止同步导致震颤增强有关。