Waxman S G, Black J A, Duncan I D, Ransom B R
Department of Neurology, Yale University School of Medicine, New Haven, Connecticut 06510.
J Neurocytol. 1990 Feb;19(1):11-28. doi: 10.1007/BF01188436.
The macromolecular structure of the axon membrane in optic nerves from 25-day-old male littermate control and myelin deficient (md) rats and 16-month-old md heterozygotic rats was examined with quantitative freeze-fracture electron microscopy. The axon membrane of control optic nerves displayed an asymmetrical partitioning of intramembranous particles (IMPs); P-fracture faces of myelinated internodal axon membrane were more particulate than those of pre-myelinated axons (approximately 1600 v 1100 microns-2, respectively), while relatively few IMPs (approximately 150 microns-2) were present on external faces (E-faces) of internodal or pre-myelinated axon membrane. Amyelinated axons of md optic nerves also exhibited an asymmetrical partitioning of IMPs; protoplasmic membrane face (P-face) IMP densities, taken as a group, exhibited a wide range (approximately 600-2300 microns-2) and, in most regions, E-faces displayed a relatively low IMP density (approximately 175 microns-2). Axons of greater than 0.4 microns diameter exhibited significantly greater mean P-face IMP density than axons less than 0.4 microns diameter. Aggregations of E-face IMPs (approximately 350 microns-2) were occasionally observed along amyelinated axon membrane from md optic nerves. Optic nerves from md heterozygote rats exhibit myelin mosaicism, permitting examination of myelinated and amyelinated axon membrane along the same tract. The axon membrane exhibits different ultrastructure in these two domains. Myelinated internodal axon membrane from md heterozygote optic nerves exhibits similar P- and E-face IMP densities to those of control internodal axolemma (approximately 1800 and 140 microns-2, respectively). Amyelinated axons in the heterozygote exhibit a membrane structure similar to amyelinated axons in md optic nerve. P-face IMP density of large diameter (greater than 0.4 microns) amyelinated axons from md heterozygote optic nerves is significantly greater than that of small calibre (less than 0.4 microns) axons. In most regions, amyelinated axon membrane exhibits a relatively low E-face IMP density (approximately 200 microns-2); however, focal aggregations (approximately 400 microns-2) of E-face particles are present. Electrophysiological recordings demonstrate that amyelinated axons in md optic nerves support the conduction of action potentials. Compound action potentials in md optic nerves exhibit a monophasic configuration, even at 20-days postnatal, similar to that of pre-myelinated optic nerve of 7-day-old normal rats. Moreover, conduction velocities in the amyelinated 20-day-old md optic nerve are similar to those displayed by pre-myelinated axons from 7-day-old optic nerves.(ABSTRACT TRUNCATED AT 400 WORDS)
采用定量冷冻断裂电子显微镜技术,研究了25日龄雄性同窝对照大鼠、髓磷脂缺乏(md)大鼠以及16月龄md杂合子大鼠视神经轴突膜的大分子结构。对照视神经的轴突膜显示出膜内颗粒(IMPs)的不对称分布;有髓鞘的节间轴突膜的P面断裂面比髓鞘形成前的轴突膜含有更多颗粒(分别约为1600和1100个/μm²),而节间或髓鞘形成前轴突膜的外表面(E面)上存在相对较少的IMPs(约150个/μm²)。md视神经的无髓鞘轴突也表现出IMPs的不对称分布;作为一个整体,原生质膜面(P面)的IMP密度范围很广(约600 - 2300个/μm²),并且在大多数区域,E面显示出相对较低的IMP密度(约175个/μm²)。直径大于0.4μm的轴突比直径小于0.4μm的轴突表现出显著更高的平均P面IMP密度。偶尔会在md视神经的无髓鞘轴突膜上观察到E面IMP的聚集(约350个/μm²)。md杂合子大鼠的视神经表现出髓鞘镶嵌现象,从而能够沿着同一条神经束检查有髓鞘和无髓鞘的轴突膜。在这两个区域,轴突膜表现出不同的超微结构。md杂合子视神经中有髓鞘的节间轴突膜表现出与对照节间轴膜相似的P面和E面IMP密度(分别约为1800和140个/μm²)。杂合子中的无髓鞘轴突表现出与md视神经中的无髓鞘轴突相似的膜结构。md杂合子视神经中直径较大(大于0.4μm)的无髓鞘轴突的P面IMP密度显著高于小口径(小于0.4μm)轴突。在大多数区域,无髓鞘轴突膜表现出相对较低的E面IMP密度(约200个/μm²);然而,存在E面颗粒的局部聚集(约400个/μm²)。电生理记录表明,md视神经中的无髓鞘轴突支持动作电位的传导。md视神经中的复合动作电位呈现单相构型,即使在出生后20天也是如此,类似于7日龄正常大鼠的髓鞘形成前视神经。此外,20日龄md视神经中无髓鞘轴突的传导速度与7日龄视神经中髓鞘形成前轴突的传导速度相似。(摘要截断于400字)