Anderson M J, Waxman S G
J Hirnforsch. 1983;24(4):371-98.
This review discusses regeneration of spinal neurons in inframammalian vertebrates. In contrast to the limited regeneration of nerve fibers in mammalian spinal cord, relatively complete regeneration of spinal cord, including the formation of new neurons, occurs in cyclostomes, selachians, teleosts, and urodele amphibians. The various experimental techniques which have been used to accelerate or enhance regeneration of spinal cord, such as implantation of peripheral nerve bridges, application of growth factors or chemical agents, and the application of exogenous electrical fields, are discussed. Processes involved in regeneration of other systems, e.g., the trophic effect of nerve fibers, the effect of certain hormones, and the effects of the blastema, are discussed as they may relate to regeneration in the spinal cord. Recent work on the factors which influence nerve outgrowth is discussed. Factors such as diffusible chemical gradients, contact with other cells, pre-formed channels for nerve growth, and differences in adhesivity of neurites for the substratum may act to control the direction of axon outgrowth in regeneration as well as normal embryonic development. This article next reviews work from our laboratory on regeneration of spinal neurons in the teleost, Sternarchus albifrons. We have used Sternarchus as a model system for the study of regeneration and remyelination since: 1. spinal cord readily regenerates after amputation of the tail, 2. new nerve cell bodies are formed during this regeneration, and 3. Sternarchus contains large neurogenic electric organs which also regenerate after tail amputation. The axons of the electric organ arise from spinal neurons and have a highly specific and complex morphology which is recapitulated after regeneration. Studies on Sternarchus indicate that new nerve cell bodies are produced during regeneration from the enlarged ependymal cells. Initially, many more of the electromotor neurons are produced in regenerated spinal cord than are present normally. Cell death subsequently occurs in some of these asynaptic cells, reducing the number of electrocytes in older regenerated cord to a more normal condition. The spinal electromotor neurons in Sternarchus have also been used to demonstrate that section of the axon in an asynaptic neuron will produce a typical "axon reaction" in the cell body. It is interesting that cell death during regeneration and the retrograde axon reaction both appear to occur without loss of synaptic elements in these normally asynaptic cells. Neurites grow through regenerating Sternarchus spinal cord in small bundles, in close contact with adjacent ependymal cell processes. Longitudinal channels, filled with neurites, are seen extending through many stages of regenerating Sternarchus cord, usually near the periphery of the cord. There are numerous invaginations of the peripheral edge of regenerating cord and its associated external lamina. These infoldings may be involved in the generation of the neurite filled channels.(ABSTRACT TRUNCATED AT 400 WORDS)
本综述讨论了非哺乳类脊椎动物脊髓神经元的再生。与哺乳动物脊髓神经纤维的有限再生不同,圆口纲动物、鲨鱼、硬骨鱼和有尾两栖动物的脊髓能发生相对完全的再生,包括新神经元的形成。文中讨论了用于加速或增强脊髓再生的各种实验技术,如植入外周神经桥、应用生长因子或化学试剂以及施加外源电场。还讨论了其他系统再生过程中涉及的相关进程,如神经纤维的营养作用、某些激素的作用以及芽基的作用,因为它们可能与脊髓再生有关。文中还讨论了近期关于影响神经生长的因素的研究。诸如可扩散化学梯度、与其他细胞的接触、预先形成的神经生长通道以及神经突与基质黏附性差异等因素,可能在再生以及正常胚胎发育过程中控制轴突生长的方向。本文接下来综述了我们实验室关于硬骨鱼白线电鳗脊髓神经元再生的研究工作。我们将白线电鳗用作研究再生和髓鞘再生的模型系统,原因如下:1. 尾巴截断后脊髓能轻易再生;2. 再生过程中会形成新的神经细胞体;3. 白线电鳗含有大型神经源性电器官,尾巴截断后该器官也能再生。电器官的轴突源自脊髓神经元,具有高度特异且复杂的形态,再生后会重现这种形态。对白线电鳗的研究表明,再生过程中扩大的室管膜细胞会产生新的神经细胞体。最初,再生脊髓中产生的电运动神经元比正常情况下的数量更多。随后,这些无突触细胞中的一些会发生细胞死亡,使较老的再生脊髓中的电细胞数量减少到更正常的状态。白线电鳗的脊髓电运动神经元还被用于证明,无突触神经元的轴突切断会在细胞体中产生典型的“轴突反应”。有趣的是,在这些通常无突触的细胞中,再生过程中的细胞死亡和逆行轴突反应似乎都是在突触元件未丢失的情况下发生的。神经突以小束状穿过再生的白线电鳗脊髓,与相邻的室管膜细胞突起紧密接触。可以看到充满神经突的纵向通道贯穿白线电鳗脊髓再生的多个阶段,通常靠近脊髓周边。再生脊髓的周边边缘及其相关的外膜有许多内陷。这些褶皱可能参与了充满神经突的通道的形成。(摘要截选至400词)