Strehler B L
University of Southern California, Los Angeles 90089.
Synapse. 1990;5(1):1-32. doi: 10.1002/syn.890050102.
A new theory designed to explain the functions of the cerebellum in the control of movement and the unique anatomy of that structure is presented. The heart of this proposed explanation is that the cerebellum generates increased numbers of outputs from particular Purkinje cells whenever the same pattern of pulses in time is presented both to its mossy fiber and its climbing fiber input systems. The postulated function of the unusual anatomy of the cerebellum is to permit it instantly to recognize and respond to identical patterns presented through these two channels regardless of the phase differences between these two signal sources, where phase differences are defined as differences in times of arrival of patterns of inputs from these two sources. The first means putatively used involves the summation of pulses comprising a given pattern of inputs simultaneously at many different Purkinje cells by virtue of the different speeds of conduction of the parallel fiber axons of granule cells. The second means is the addition of an input from the climbing fiber system that, together with the simultaneous parallel fiber inputs, leads to a discharge of particular Purkinje cells, which discharge temporarily increases the size of EPSP's generated by the parallel fiber synapses involved in the cell's discharge. This specific synaptic potentiation, in turn, makes it possible for the cell to respond by generating closely consecutive additional discharges provided that the same patterns of discharge are presented both to the climbing fiber system and the mossy fiber system. This happens because later pulses in identical patterns will arrive simultaneously at previously facilitated synapses via parallel fibers and at synapses of the climbing fibers, thereby causing additional spatial summations and discharges to occur. According to this explanation the patterns that are compared in the above manner are symbols (patterns of pulses) produced by sensors of current positions and symbols derived from memory and also representing these same positions. When patterns from these two sources are identical, the multiple outputs of specific Purkinje cells inhibit an automatic feedback loop and thereby indirectly cause the attenuation and arrest of movement. The evolution of these concepts resulted in very specific "predictions" of particular connections involving the olive, pons, red nucleus, dentate, thalamus, and sensory-motor cortex. All of these predictions were found to be consistent with evidence in the literature. Points of difference between this theory and all prior ones are discussed as are several critical tests of its validity, and the putative evolution of cerebellar structure and function.(ABSTRACT TRUNCATED AT 400 WORDS)
本文提出了一种新理论,旨在解释小脑在运动控制中的功能及其独特的解剖结构。该理论的核心观点是,每当相同的脉冲时间模式同时呈现给小脑的苔藓纤维和攀缘纤维输入系统时,小脑特定浦肯野细胞的输出数量就会增加。小脑独特解剖结构的假定功能是使其能够立即识别并响应通过这两个通道呈现的相同模式,而不管这两个信号源之间的相位差异如何,相位差异定义为来自这两个源的输入模式到达时间的差异。第一种可能的方式涉及颗粒细胞平行纤维轴突传导速度不同,从而在许多不同的浦肯野细胞中同时对构成给定输入模式的脉冲进行求和。第二种方式是来自攀缘纤维系统的输入与同时的平行纤维输入相加,导致特定浦肯野细胞放电,这种放电会暂时增加该细胞放电所涉及的平行纤维突触产生的兴奋性突触后电位(EPSP)的大小。这种特定的突触增强反过来又使细胞能够通过产生紧密连续的额外放电做出反应,前提是相同的放电模式同时呈现给攀缘纤维系统和苔藓纤维系统。这是因为相同模式中的后续脉冲将同时通过平行纤维到达先前易化的突触以及攀缘纤维的突触,从而导致额外的空间总和与放电发生。根据这种解释,以上述方式进行比较的模式是由当前位置传感器产生的符号(脉冲模式)以及来自记忆且也代表相同位置的符号。当来自这两个源的模式相同时,特定浦肯野细胞的多个输出会抑制自动反馈回路,从而间接导致运动的减弱和停止。这些概念的演变导致了对涉及橄榄核、脑桥、红核、齿状核、丘脑和感觉运动皮层的特定连接的非常具体的“预测”。所有这些预测都被发现与文献中的证据一致。本文讨论了该理论与所有先前理论的不同之处,以及对其有效性的几个关键测试,还有小脑结构和功能的假定演变。(摘要截选至400字)