Hof P R, Glezer I I, Condé F, Flagg R A, Rubin M B, Nimchinsky E A, Vogt Weisenhorn D M
Fishberg Research Center for Neurobiology, Department of Geriatrics and Adult Development, Mount Sinai School of Medicine, New York, NY 10029, USA.
J Chem Neuroanat. 1999 Feb;16(2):77-116. doi: 10.1016/s0891-0618(98)00065-9.
The three calcium-binding proteins parvalbumin, calbindin, and calretinin are found in morphologically distinct classes of inhibitory interneurons as well as in some pyramidal neurons in the mammalian neocortex. Although there is a wide variability in the qualitative and quantitative characteristics of the neocortical subpopulations of calcium-binding protein-immunoreactive neurons in mammals, most of the available data show that there is a fundamental similarity among the mammalian species investigated so far, in terms of the distribution of parvalbumin, calbindin, and calretinin across the depth of the neocortex. Thus, calbindin- and calretinin-immunoreactive neurons are predominant in layers II and III, but are present across all cortical layers, whereas parvalbumin-immunoreactive neurons are more prevalent in the middle and lower cortical layers. These different neuronal populations have well defined regional and laminar distribution, neurochemical characteristics and synaptic connections, and each of these cell types displays a particular developmental sequence. Most of the available data on the development, distribution and morphological characteristics of these calcium-binding proteins are from studies in common laboratory animals such as the rat, mouse, cat, macaque monkey, as well as from postmortem analyses in humans, but there are virtually no data on other species aside of a few incidental reports. In the context of the evolution of mammalian neocortex, the distribution and morphological characteristics of calcium-binding protein-immunoreactive neurons may help defining taxon-specific patterns that may be used as reliable phylogenetic traits. It would be interesting to extend such neurochemical analyses of neuronal subpopulations to other species to assess the degree to which neurochemical specialization of particular neuronal subtypes, as well as their regional and laminar distribution in the cerebral cortex, may represent sets of derived features in any given mammalian order. This could be particularly interesting in view of the consistent differences in neurochemical typology observed in considerably divergent orders such as cetaceans and certain families of insectivores and metatherians, as well as in monotremes. The present article provides an overview of calcium-binding protein distribution across a large number of representative mammalian species and a review of their developmental patterns in the species where data are available. This analysis demonstrates that while it is likely that the developmental patterns are quite consistent across species, at least based on the limited number of species for which ontogenetic data exist, the distribution and morphology of calcium-binding protein-containingneurons varies substantially among mammalian orders and that certain species show highly divergent patterns compared to closely related taxa. Interestingly, primates, carnivores, rodents and tree shrews appear closely related on the basis of the observed patterns, marsupials show some affinities with that group, whereas prototherians have unique patterns. Our findings also support the relationships of cetaceans and ungulates, and demonstrates possible affinities between carnivores and ungulates, as well as the existence of common, probably primitive, traits in cetaceans and insectivores.
三种钙结合蛋白——小白蛋白、钙结合蛋白和钙视网膜蛋白,存在于哺乳动物新皮层中形态各异的抑制性中间神经元类别以及一些锥体神经元中。尽管哺乳动物新皮层中钙结合蛋白免疫反应性神经元亚群的定性和定量特征存在很大差异,但大多数现有数据表明,就小白蛋白、钙结合蛋白和钙视网膜蛋白在新皮层各深度的分布而言,到目前为止所研究的哺乳动物物种之间存在基本相似性。因此,钙结合蛋白和钙视网膜蛋白免疫反应性神经元在第II层和第III层中占主导地位,但在所有皮层层中都有分布,而小白蛋白免疫反应性神经元在皮层中层和下层更为普遍。这些不同的神经元群体具有明确的区域和层状分布、神经化学特征和突触连接,并且每种细胞类型都表现出特定的发育序列。关于这些钙结合蛋白的发育、分布和形态特征的大多数现有数据来自对常见实验动物(如大鼠、小鼠、猫、猕猴)的研究,以及对人类的尸检分析,但除了一些偶然的报告外,几乎没有关于其他物种的数据。在哺乳动物新皮层进化的背景下,钙结合蛋白免疫反应性神经元的分布和形态特征可能有助于确定特定分类群的模式,这些模式可作为可靠的系统发育特征。将这种对神经元亚群的神经化学分析扩展到其他物种,以评估特定神经元亚型的神经化学特化程度,以及它们在大脑皮层中的区域和层状分布在任何给定的哺乳动物目中可能代表衍生特征集的程度将是很有趣的。鉴于在鲸类、某些食虫动物和有袋动物科以及单孔目等差异较大的目中观察到的神经化学类型存在一致差异,这可能会特别有趣。本文概述了大量代表性哺乳动物物种中钙结合蛋白的分布,并回顾了在有数据的物种中的发育模式。该分析表明,虽然发育模式可能在物种间相当一致,至少基于存在个体发育数据的有限物种数量来看是这样,但含钙结合蛋白神经元的分布和形态在哺乳动物目中有很大差异,并且某些物种与密切相关的分类群相比表现出高度不同的模式。有趣的是,根据观察到的模式,灵长类动物、食肉动物、啮齿动物和树鼩似乎关系密切,有袋动物与该群体有一些亲缘关系,而原兽亚纲动物有独特的模式。我们的研究结果也支持鲸类和有蹄类动物的关系,并证明了食肉动物和有蹄类动物之间可能的亲缘关系,以及鲸类和食虫动物中存在共同的、可能是原始的特征。