Kolb F P, Arnold G, Lerch R, Straka H, Büttner-Ennever J
Institute of Physiology, University of Munich, Germany.
Neuroscience. 1997 Dec;81(4):1155-81. doi: 10.1016/s0306-4522(97)00255-8.
The present study was designed to characterize the spread of excitation within the frontal plane of the cat cerebellar cortex following different types of stimuli. In particular, experiments were performed to determine whether the spread of excitation evoked by mossy fibre inputs proceeds primarily along the parallel fibres ("beam-like" spread) or whether these inputs activate non-propagated foci ("patches") in the cerebellar cortex. Field potentials were recorded within a frontal plane as a medial to lateral array at different depths in parallel tracks. The recordings were made following electrical stimulation of different forelimb nerves and functionally related areas of the sensorimotor cortex as well as during passive paw movements. The resulting spatial grid of responses provides discrete spatio-temporal information reflecting the activation of specific cerebellar afferents and the neuronal interactions they evoke. The method employed demonstrates the spatial distribution of the temporal sequence of excitability changes throughout all the cerebellar cortical layers. In general, the characteristics of the responses in the intermediate cerebellar cortex depended on the source of the signals. Activity patterns evoked by peripheral nerve stimulation showed more clustered foci compared with those following electrical stimulation of functionally related areas of the sensorimotor cortex. The centrally evoked profiles were generally more homogeneous. The largest number of foci were observed following passive movements around the wrist joint. The spread of excitation in the vertical direction was evaluated by the spatial shift of the line of reversal of the N3/P2-potential (zero-isopotential line). Lines of reversal for peripherally-evoked activity patterns were approximately 90 microns closer to the molecular layer than those evoked by central stimulation in animals in which recordings have been performed in lobule Vc. The opposite was found for recordings in lobule Vb, where potential reversals following peripheral stimulation were located 40 microns deeper than those evoked following central stimulation. Cortical inputs resulted in a more proximal activation of lobule Vc Purkinje cell dendrites than in lobule Vb. This type of input processing thus seems to be lobule dependent. A beam-like spread of excitation could not be demonstrated. For both climbing fibre and mossy fibre afferent systems multiple foci were found in the frontal plane. The foci due to mossy fibre activation arose from the granular layer and expanded vertically to the molecular layer. For the climbing fibre system the foci were restricted to the molecular layer, where they merged to form a superficial band of activation. Although the data presented in this paper favour a focal distribution of activity, they do not exclude beam-like propagation along the parallel fibres, because of the difficulty of detecting this pattern in response to the stimuli. The "beam"- and "patch"-like hypotheses need not be mutually exclusive. Each could contribute to a specific stage of the temporal-spatial processing in the cerebellar cortex in a functional and task-specific manner.
本研究旨在描述猫小脑皮质额面内不同类型刺激后兴奋的传播特征。具体而言,进行实验以确定苔藓纤维输入所诱发的兴奋传播是否主要沿着平行纤维进行(“束状”传播),或者这些输入是否激活小脑皮质中的非传播性焦点(“斑块”)。在额面内,沿着平行轨道在不同深度以从内侧到外侧的阵列记录场电位。记录是在电刺激不同的前肢神经和感觉运动皮质的功能相关区域后以及在被动爪子运动期间进行的。由此产生的空间反应网格提供了离散的时空信息,反映了特定小脑传入神经的激活及其引发的神经元相互作用。所采用的方法展示了兴奋性变化的时间序列在所有小脑皮质层中的空间分布。一般来说,小脑中间皮质的反应特征取决于信号来源。与感觉运动皮质功能相关区域的电刺激相比,外周神经刺激诱发的活动模式显示出更多聚集的焦点。中枢诱发的轮廓通常更均匀。在腕关节周围的被动运动后观察到的焦点数量最多。通过N3/P2电位反转线(零等电位线)的空间移位来评估垂直方向上的兴奋传播。在小叶Vc进行记录的动物中,外周诱发活动模式的反转线比中枢刺激诱发的反转线更靠近分子层约90微米。在小叶Vb的记录中发现了相反的情况,外周刺激后的电位反转比中枢刺激后的电位反转深40微米。皮质输入导致小叶Vc浦肯野细胞树突的激活比小叶Vb更靠近近端。因此,这种类型的输入处理似乎依赖于小叶。无法证明兴奋的束状传播。对于攀缘纤维和苔藓纤维传入系统,在额面中发现了多个焦点。苔藓纤维激活产生的焦点起源于颗粒层,并垂直扩展到分子层。对于攀缘纤维系统,焦点局限于分子层,在那里它们合并形成一个浅表的激活带。尽管本文给出的数据支持活动的焦点分布,但由于难以检测到对刺激的这种模式,它们并不排除沿着平行纤维的束状传播。“束状”和“斑块状”假说不一定相互排斥。每种假说都可以以功能和任务特定的方式对小脑皮质时空处理的特定阶段做出贡献。