Faculty of Biology, Medicine and Health, The University of Manchester, Manchester, M13 9PT, UK.
Faculty of Health and Medical Sciences, Center for Basic and Translational Neuroscience, University of Copenhagen, 2200, Copenhagen, Denmark.
Adv Exp Med Biol. 2019;1175:15-44. doi: 10.1007/978-981-13-9913-8_2.
As the nervous system evolved from the diffused to centralised form, the neurones were joined by the appearance of the supportive cells, the neuroglia. Arguably, these non-neuronal cells evolve into a more diversified cell family than the neurones are. The first ancestral neuroglia appeared in flatworms being mesenchymal in origin. In the nematode C. elegans proto-astrocytes/supportive glia of ectodermal origin emerged, albeit the ensheathment of axons by glial cells occurred later in prawns. The multilayered myelin occurred by convergent evolution of oligodendrocytes and Schwann cells in vertebrates above the jawless fishes. Nutritive partitioning of the brain from the rest of the body appeared in insects when the hemolymph-brain barrier, a predecessor of the blood-brain barrier was formed. The defensive cellular mechanism required specialisation of bona fide immune cells, microglia, a process that occurred in the nervous system of leeches, bivalves, snails, insects and above. In ascending phylogeny, new type of glial cells, such as scaffolding radial glia, appeared and as the bran sizes enlarged, the glia to neurone ratio increased. Humans possess some unique glial cells not seen in other animals.
随着神经系统从弥散形式进化为集中形式,神经元出现了支持细胞,即神经胶质。可以说,这些非神经元细胞比神经元进化成了更加多样化的细胞家族。第一批祖先进化而来的神经胶质出现在扁形动物中,起源于间质。在线虫 C. elegans 中,出现了源自外胚层的原星形胶质细胞/支持胶质,但轴突被胶质细胞包裹的现象在虾类中出现得较晚。多层髓鞘是由无颌鱼类以上的少突胶质细胞和施万细胞的趋同进化形成的。当血液-脑屏障的前身血淋巴-脑屏障形成时,昆虫的大脑与身体其他部分的营养分配出现了分化。在防御细胞机制方面,需要专门化的真正免疫细胞,即小胶质细胞,这一过程发生在水蛭、双壳类、蜗牛、昆虫及以上动物的神经系统中。随着系统发生的上升,新类型的神经胶质细胞,如支架状放射状胶质细胞,出现了,而且随着脑区的增大,神经胶质细胞与神经元的比例增加了。人类拥有一些在其他动物中没有的独特的神经胶质细胞。