Wenzel J, Bogolepov N N
J Hirnforsch. 1976;17(5):399-448.
The neuronal structure and synaptology of the hippocampus of the rat are studied using neurohistological, electron microscopical and morphometrical methods. After Golgi-impregnations the neurons of the hippocampus were classified by reason of special morphological features (pericaryon, dendritic, ramification, course and termination of the axons). The neurons of the hippocampus show from the region CA4 towards the region CA1 an increasing "pyramidalization" which is explained respecting the neuronal structure and synaptic architectonics as the result of a specific presynaptic representation of afferents at the neurons. The high degree of differentiation of the CA1-pyramids speaks for the progressive developmental tendency in the phylogenesis of these pyramids compared to the neocortical neurons. The ultrastructural investigations of the hippocampus show that each individual layer of the regions CA1, CA3 and CA4 has a characteristic synaptic architectonics. In all hippocampal layers the axo-spino-dendritic contacts are the most frequent type of synapses with the structural characteristics of the excitatory synapses; the synaptic contact appears as "Bouton en passage" respectively as "Bouton terminal". The axosomatic synapses of the stratum pyramidale (CA1 to CA3) predominantly are developed as symmetrical contacts with inhibitory function. As the terminals of the basket cells they represent the morphological correlate for the recurrent inhibitation of the pyramids by the inhibitory effecting basket cells. In the supra- and infrapyramidal layers of the regions CA3 and CA4 special contacts are developed for the connection withe the mossy fiber terminals--at the pericarya and the main dendrites as so-called somatic and dendritic protrusions and as micro-dendrites. As has been established for the neocortex, the dendritic spines are the decisive receptive postsynaptic structural characteristic of the hippocampus as well. Pre- and postsynaptic elements show a broad scope of variation in forming their structures. This also applies to the intrahippocampal systems with their highly complex structure which is the morphological expression of an extremely high degree of differentiation. The morphometrical investigations of the ultrastructure of synapses enclose the calculation of the number of synapses per mm3 of the volume of layers, the average individual volume of synapses, the presynaptic portion of volume of the relative volume of cortex of the hippocampal layers as well as the average number and size of the synaptic vesicles. The average number of synapses per mm3 brain tissue runs in CA1 to 5,9 X 10(9), in CA3 to 7,9 X 10(9) andin CA4 to 8,7 X 10(9) synapses per mm3. The average individual volume of synapses amounts to 0,2-0,3 mm3 whereas the average portion of volume of the presynaptic structures per relative cortical volume comes to 14% in CA1, 16-19% in CA3 and 15% in CA4.
采用神经组织学、电子显微镜和形态计量学方法研究大鼠海马的神经元结构和突触学。经高尔基氏染色后,根据海马神经元特殊的形态学特征(胞体、树突、分支、轴突的走行和终末)进行分类。海马神经元从CA4区到CA1区呈现出逐渐增强的“锥化”现象,从神经元结构和突触构筑学角度来看,这是传入神经元在这些神经元处特定的突触前表现的结果。CA1区锥体神经元的高度分化表明,与新皮质神经元相比,这些锥体神经元在系统发生过程中具有渐进的发育趋势。海马的超微结构研究表明,CA1、CA3和CA4区的每一层都有其独特的突触构筑。在所有海马层中,轴-棘-树突接触是最常见的突触类型,具有兴奋性突触的结构特征;突触接触分别表现为“中途扣结”或“终末扣结”。锥体层(CA1至CA3)中的轴-体突触主要发育为具有抑制功能的对称接触。作为篮状细胞的终末,它们代表了抑制性篮状细胞对锥体神经元进行回返抑制的形态学关联。在CA3和CA4区的锥体上、下层,为与苔藓纤维终末建立联系而形成了特殊的接触——在胞体和主要树突上,表现为所谓的体部和树突状突起以及微树突。正如在新皮质中所证实的那样,树突棘也是海马决定性的突触后接受结构特征。突触前和突触后成分在形成其结构时表现出广泛的变化。这也适用于具有高度复杂结构的海马内系统,这种复杂结构是高度分化程度的形态学表现。对突触超微结构的形态计量学研究包括计算每立方毫米各层体积中的突触数量、突触的平均个体体积、海马层皮质相对体积中突触前结构的体积比例以及突触小泡的平均数量和大小。每立方毫米脑组织中的突触平均数量在CA1区为5.9×10⁹个,在CA3区为7.9×10⁹个,在CA4区为8.7×10⁹个。突触的平均个体体积为0.2 - 0.3立方毫米,而相对于皮质体积,突触前结构的平均体积比例在CA1区为14%,在CA3区为16 - 19%,在CA4区为15%。