Hellwig B
Max-Planck-Institute für biologische Kybernetik, Tübingen, Germany.
J Hirnforsch. 1993;34(3):387-402.
In the cerebral cortex two main types of anatomical parcellation have been proposed: cyto- and myeloarchitectonics. Cytoarchitectonics is based on differences in the distribution and the sizes of cell bodies. Myeloarchitectonics relies on the layering and packing density of intracortical myelinated fibres. Thereby attention is mainly focused on those horizontal fibres which are organized in bands known as the stripes of Baillarger. Cyto- and myeloarchitectonics must be somehow related: structural changes from area to area as revealed by the Nissl stain coincide with changes in the myelin picture. In this paper, it is demonstrated that two simple assumptions are sufficient to transform quantitative data on the Nissl picture of a certain area (i.e. layer thicknesses, neuron sizes, neuron densities) into the corresponding myelin picture. The first assumption is that large neurons contribute more to the intracortical myelin content than small ones, and that this relation can be represented by a sigmoid curve. The second assumption is that the average distribution of horizontal axon collaterals of pyramidal neurons with respect to the cell body can be quantified by a simple model. On the basis of these two assumptions, myelin patterns were computed for the whole spectrum of cortical variability, including the primary visual, somatosensory, auditory and motor cortices, the speech centres and a number of association areas. Comparison of the simulations with real myelin preparations revealed remarkable similarities. These findings support the assumption that the horizontal component of the myelin picture is mainly produced by axon collaterals of pyramidal cells, and that nonpyramidal neurons and afferent fibres play a minor role. Moreover, the results suggest that the distribution of horizontal axon collaterals of pyramidal neurons and the principles of their myelination are very similar in different areas. The function of intracortical myelin is discussed. The increase in conduction velocity gained by myelination seems negligible for most intracortical fibres. It is argued that myelination may be related to learning processes during the critical period.
在大脑皮层中,已提出两种主要的解剖学分区类型:细胞构筑学和髓鞘构筑学。细胞构筑学基于细胞体分布和大小的差异。髓鞘构筑学则依赖于皮质内有髓纤维的分层和堆积密度。因此,注意力主要集中在那些以贝亚尔热带(Baillarger条纹)形式成带排列的水平纤维上。细胞构筑学和髓鞘构筑学必定在某种程度上相关:尼氏染色所揭示的不同区域间的结构变化与髓鞘图像的变化相一致。在本文中,证明了两个简单假设足以将某一区域尼氏图像的定量数据(即层厚度、神经元大小、神经元密度)转换为相应的髓鞘图像。第一个假设是,大神经元比小神经元对皮质内髓鞘含量的贡献更大,且这种关系可用S形曲线表示。第二个假设是,锥体神经元水平轴突侧支相对于细胞体的平均分布可用一个简单模型进行量化。基于这两个假设,计算了包括初级视觉、躯体感觉、听觉和运动皮层、言语中枢以及一些联合区域在内的整个皮质变异性范围内的髓鞘模式。模拟结果与真实髓鞘标本的比较显示出显著的相似性。这些发现支持了这样一种假设,即髓鞘图像的水平成分主要由锥体细胞的轴突侧支产生,而非锥体神经元和传入纤维起次要作用。此外,结果表明锥体神经元水平轴突侧支的分布及其髓鞘形成原则在不同区域非常相似。文中还讨论了皮质内髓鞘的功能。髓鞘化所带来的传导速度增加对大多数皮质内纤维而言似乎微不足道。有人认为髓鞘化可能与关键期的学习过程有关。