Wu Di, Klaw Michelle C, Kholodilov Nikolai, Burke Robert E, Detloff Megan R, Côté Marie-Pascale, Tom Veronica J
Department of Neurobiology and Anatomy, Drexel University College of Medicine Philadelphia, PA, USA.
Department of Neurology, Columbia University in the City of New York New York, NY, USA.
Front Mol Neurosci. 2016 Jul 5;9:49. doi: 10.3389/fnmol.2016.00049. eCollection 2016.
While the peripheral branch of dorsal root ganglion neurons (DRG) can successfully regenerate after injury, lesioned central branch axons fail to regrow across the dorsal root entry zone (DREZ), the interface between the dorsal root and the spinal cord. This lack of regeneration is due to the limited regenerative capacity of adult sensory axons and the growth-inhibitory environment at the DREZ, which is similar to that found in the glial scar after a central nervous system (CNS) injury. We hypothesized that transduction of adult DRG neurons using adeno-associated virus (AAV) to express a constitutively-active form of the GTPase Rheb (caRheb) will increase their intrinsic growth potential after a dorsal root crush. Additionally, we posited that if we combined that approach with digestion of upregulated chondroitin sulfate proteoglycans (CSPG) at the DREZ with chondroitinase ABC (ChABC), we would promote regeneration of sensory axons across the DREZ into the spinal cord. We first assessed if this strategy promotes neuritic growth in an in vitro model of the glial scar containing CSPG. ChABC allowed for some regeneration across the once potently inhibitory substrate. Combining ChABC treatment with expression of caRheb in DRG significantly improved this growth. We then determined if this combination strategy also enhanced regeneration through the DREZ after dorsal root crush in adult rats in vivo. After unilaterally crushing C4-T1 dorsal roots, we injected AAV5-caRheb or AAV5-GFP into the ipsilateral C5-C8 DRGs. ChABC or PBS was injected into the ipsilateral dorsal horn at C5-C8 to digest CSPG, for a total of four animal groups (caRheb + ChABC, caRheb + PBS, GFP + ChABC, GFP + PBS). Regeneration was rarely observed in PBS-treated animals, whereas short-distance regrowth across the DREZ was observed in ChABC-treated animals. No difference in axon number or length between the ChABC groups was observed, which may be related to intraganglionic inflammation induced by the injection. ChABC-mediated regeneration is functional, as stimulation of ipsilateral median and ulnar nerves induced neuronal c-Fos expression in deafferented dorsal horn in both ChABC groups. Interestingly, caRheb + ChABC animals had significantly more c-Fos(+) nuclei indicating that caRheb expression in DRGs promoted functional synaptogenesis of their axons that regenerated beyond a ChABC-treated DREZ.
虽然背根神经节神经元(DRG)的外周分支在损伤后能够成功再生,但受损的中枢分支轴突无法穿过背根进入区(DREZ),即背根与脊髓之间的界面。这种再生缺失是由于成体感觉轴突的再生能力有限以及DREZ处的生长抑制环境,该环境与中枢神经系统(CNS)损伤后胶质瘢痕中的环境相似。我们假设,使用腺相关病毒(AAV)转导成年DRG神经元以表达GTP酶Rheb的组成型活性形式(caRheb),将增加其在背根挤压后的内在生长潜力。此外,我们推测,如果将该方法与用软骨素酶ABC(ChABC)消化DREZ处上调的硫酸软骨素蛋白聚糖(CSPG)相结合,我们将促进感觉轴突穿过DREZ向脊髓的再生。我们首先在含有CSPG的胶质瘢痕体外模型中评估该策略是否促进神经突生长。ChABC使轴突能够穿过曾经具有强烈抑制作用的底物实现一定程度的再生。将ChABC处理与DRG中caRheb的表达相结合可显著改善这种生长。然后我们确定这种联合策略在成年大鼠体内背根挤压后是否也能增强穿过DREZ的再生。在单侧挤压C4 - T1背根后,我们将AAV5 - caRheb或AAV5 - GFP注入同侧C5 - C8 DRG。将ChABC或PBS注入同侧C5 - C8背角以消化CSPG,共分为四个动物组(caRheb + ChABC、caRheb + PBS、GFP + ChABC、GFP + PBS)。在PBS处理的动物中很少观察到再生,而在ChABC处理的动物中观察到穿过DREZ的短距离再生。在ChABC组之间未观察到轴突数量或长度的差异,这可能与注射引起的神经节内炎症有关。ChABC介导的再生是有功能的,因为刺激同侧正中神经和尺神经在两个ChABC组的去传入背角中均诱导神经元c - Fos表达。有趣的是,caRheb + ChABC组动物具有明显更多的c - Fos(+)细胞核,表明DRG中caRheb的表达促进了其轴突的功能性突触形成,这些轴突再生至超过ChABC处理的DREZ。