成年斑马鱼视神经损伤后轴突再生与髓鞘再建的研究
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摘要
斑马鱼中枢神经系统拥有很强的再生能力,它的视神经在损伤之后能够很快地再生并且恢复视觉功能。然而,这种视觉功能的恢复是否需要视网膜神经节细胞(retinal ganglion cell, RGC)的新生还不是很清楚。我们对不同损伤条件下RGC细胞的数目进行了定量统计,结果表明：在视神经夹伤的情况下,几乎所有的RGC都存活；在视神经切断模式中前2周有超过90%的RGC存活,并且在损伤后第7周仍旧有75%的RGC存活。顶盖逆行标记显示了RGC轴突有惊人的再生能力,在视神经夹伤后第1周有90%以上的RGC再生到了顶盖,在切断模型中也有50%以上的RGC再生到了顶盖,切断之后的第4周达到正常的水平。此外,RGC双标与BrdU的结果都显示几乎没有新的RGC产生。因此,我们直接地证明了成年斑马鱼视神经损伤后的视觉功能恢复的主要策略是RGC细胞的存活与轴突的再生。
与神经元再生障碍不同的是,中枢神经系统的髓鞘结构可以再建。关于成年动物再髓鞘化的过程,一般认为它是发育过程中髓鞘形成的重演。然而,关于再髓鞘细胞的来源问题,现在了解的还不是很全面,一般认为是少突胶质细胞前体细胞(oligodendrocyte precursor cells, OPC)充当了主要的来源,而成熟的少突胶质细胞(oligodendrocyte, OC)则不太可能在这一过程中扮演角色。即使有人在电镜下观察到了成熟的OC可能也具有分裂的能力,但是由于到目前为止还缺乏直接的证据。成年斑马鱼视神经损伤能够造成髓鞘的丢失,之后表现出再髓鞘现象。我们观察到了炎症细胞快速侵入并帮助清理髓鞘片段,但是很少看到OC胞体的消失。相反,我们看到了OC表现出增殖的现象,表达pho-ERK1/2的olig2+细胞数目在损伤远心端明显比近心端要显著提高。我们在活体水平实时动态观察到了移植视神经上具有复杂分支的OC能够以一种非对称的方式产生新的细胞,新产生的细胞类似于前体细胞,具有很强的迁移能力。我们进一步发现炎症反应在一定程度上阻碍了OPC的迁移进入损伤位点。在损伤早期阶段,损伤位点聚集大量的NG2阳性细胞,且该时刻视神经上TNFa水平最高,从而阻止OPC迁移进入损伤位点；而当NG2与TNFa消失之后,OPC开始进入损伤位点并逐渐成熟。我们用结扎的办法延长视神经上炎症的存在时间,发现OPC进入损伤位点的时间与髓鞘恢复的时间都有所延迟。最后,髓鞘再生在损伤后第7周基本完成,虽然髓鞘的厚度没有达到正常水平,但是郎飞氏节的密度并没有降低,视觉功能也在髓鞘再生基本完成的情况下得以恢复。
Zebrafish central nervous system (CNS) possesses a strong neural regeneration ability to restore visual function completely after optic nerve injury (ONI). However, whether neurogenesis of retinal ganglion cell (RGC) contributes to functional recovery remains controversial. Our quantitative analysis of RGCs in different ONI models showed that almost all RGCs survived in optic nerve crush (ONC) model; while over90%of RGCs survived in the first2weeks with75%remaining after7weeks in optic nerve transection (ONT) model. Retrograde labeling from tectum revealed a surprising regeneration rate, with over90%and over50%of RGCs regrowing axons to tectum at the first week in ONC and ONT model respectively. In the latter one, the number of regenerative RGCs after4weeks had no significant difference from the control group. As for neurogenesis, newborn RGCs were rarely detected either by double retrograde labeling or BrdU marker. Our results have directly shown that RGC survival and axon regrowth are responsible for functional recovery after ONI in adult zebrafish.
Not like neural regeneration is not happen in mammalian, remyelination is generally happen. It is widely accepted that remyelination is the recapitulation of myelination during development. Whether mature oligodendrocytes also contribute to this procedure is still ambiguity. Here, we use the model of optic nerve injury in adult zebrafish to research remyelination during functional recovery. First, optic nerve injury induced loose myelin components and inflammation cells invasion. Second, oligodendrocytes over expressed pho-ERK1/2which indicated they began to re-enter the cell cycle. Third, oligodendrocytes whose processes still remained division in an asymmetry way, and the new born cells had a strong ability of migration. Fourth, at the first week, inflammation existing in the epicenter may play a role in repelling the migration of OPCs as NG2+astrocytes existing, with the highest TNFa level at this period; in the following weeks, OPCs were recruited into the epicenter and myelin structure was recovered, which can be delayed when inflammation was elongated by ligation of the optic nerve. Lastly, the recovery of myelin index and ranvier node, combining with the restitution of fasciculate structure, contributes to the visual functional recovery.
与神经元再生障碍不同的是,中枢神经系统的髓鞘结构可以再建。关于成年动物再髓鞘化的过程,一般认为它是发育过程中髓鞘形成的重演。然而,关于再髓鞘细胞的来源问题,现在了解的还不是很全面,一般认为是少突胶质细胞前体细胞(oligodendrocyte precursor cells, OPC)充当了主要的来源,而成熟的少突胶质细胞(oligodendrocyte, OC)则不太可能在这一过程中扮演角色。即使有人在电镜下观察到了成熟的OC可能也具有分裂的能力,但是由于到目前为止还缺乏直接的证据。成年斑马鱼视神经损伤能够造成髓鞘的丢失,之后表现出再髓鞘现象。我们观察到了炎症细胞快速侵入并帮助清理髓鞘片段,但是很少看到OC胞体的消失。相反,我们看到了OC表现出增殖的现象,表达pho-ERK1/2的olig2+细胞数目在损伤远心端明显比近心端要显著提高。我们在活体水平实时动态观察到了移植视神经上具有复杂分支的OC能够以一种非对称的方式产生新的细胞,新产生的细胞类似于前体细胞,具有很强的迁移能力。我们进一步发现炎症反应在一定程度上阻碍了OPC的迁移进入损伤位点。在损伤早期阶段,损伤位点聚集大量的NG2阳性细胞,且该时刻视神经上TNFa水平最高,从而阻止OPC迁移进入损伤位点；而当NG2与TNFa消失之后,OPC开始进入损伤位点并逐渐成熟。我们用结扎的办法延长视神经上炎症的存在时间,发现OPC进入损伤位点的时间与髓鞘恢复的时间都有所延迟。最后,髓鞘再生在损伤后第7周基本完成,虽然髓鞘的厚度没有达到正常水平,但是郎飞氏节的密度并没有降低,视觉功能也在髓鞘再生基本完成的情况下得以恢复。
Zebrafish central nervous system (CNS) possesses a strong neural regeneration ability to restore visual function completely after optic nerve injury (ONI). However, whether neurogenesis of retinal ganglion cell (RGC) contributes to functional recovery remains controversial. Our quantitative analysis of RGCs in different ONI models showed that almost all RGCs survived in optic nerve crush (ONC) model; while over90%of RGCs survived in the first2weeks with75%remaining after7weeks in optic nerve transection (ONT) model. Retrograde labeling from tectum revealed a surprising regeneration rate, with over90%and over50%of RGCs regrowing axons to tectum at the first week in ONC and ONT model respectively. In the latter one, the number of regenerative RGCs after4weeks had no significant difference from the control group. As for neurogenesis, newborn RGCs were rarely detected either by double retrograde labeling or BrdU marker. Our results have directly shown that RGC survival and axon regrowth are responsible for functional recovery after ONI in adult zebrafish.
Not like neural regeneration is not happen in mammalian, remyelination is generally happen. It is widely accepted that remyelination is the recapitulation of myelination during development. Whether mature oligodendrocytes also contribute to this procedure is still ambiguity. Here, we use the model of optic nerve injury in adult zebrafish to research remyelination during functional recovery. First, optic nerve injury induced loose myelin components and inflammation cells invasion. Second, oligodendrocytes over expressed pho-ERK1/2which indicated they began to re-enter the cell cycle. Third, oligodendrocytes whose processes still remained division in an asymmetry way, and the new born cells had a strong ability of migration. Fourth, at the first week, inflammation existing in the epicenter may play a role in repelling the migration of OPCs as NG2+astrocytes existing, with the highest TNFa level at this period; in the following weeks, OPCs were recruited into the epicenter and myelin structure was recovered, which can be delayed when inflammation was elongated by ligation of the optic nerve. Lastly, the recovery of myelin index and ranvier node, combining with the restitution of fasciculate structure, contributes to the visual functional recovery.
引文
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