Niemi Jon P, DeFrancesco-Lisowitz Alicia, Cregg Jared M, Howarth Madeline, Zigmond Richard E
Department of Neurosciences, Case Western Reserve University, Cleveland, OH 44106-4975, USA.
Department of Neurosciences, Case Western Reserve University, Cleveland, OH 44106-4975, USA.
Exp Neurol. 2016 Jan;275 Pt 1(0 1):25-37. doi: 10.1016/j.expneurol.2015.09.018. Epub 2015 Nov 2.
Neuroinflammation plays a critical role in the regeneration of peripheral nerves following axotomy. An injury to the sciatic nerve leads to significant macrophage accumulation in the L5 DRG, an effect not seen when the dorsal root is injured. We recently demonstrated that this accumulation around axotomized cell bodies is necessary for a peripheral conditioning lesion response to occur. Here we asked whether overexpression of the monocyte chemokine CCL2 specifically in DRG neurons of uninjured mice is sufficient to cause macrophage accumulation and to enhance regeneration or whether other injury-derived signals are required. AAV5-EF1α-CCL2 was injected intrathecally, and this injection led to a time-dependent increase in CCL2 mRNA expression and macrophage accumulation in L5 DRG, with a maximal response at 3 weeks post-injection. These changes led to a conditioning-like increase in neurite outgrowth in DRG explant and dissociated cell cultures. This increase in regeneration was dependent upon CCL2 acting through its primary receptor CCR2. When CCL2 was overexpressed in CCR2-/- mice, macrophage accumulation and enhanced regeneration were not observed. To address the mechanism by which CCL2 overexpression enhances regeneration, we tested for elevated expression of regeneration-associated genes in these animals. Surprisingly, we found that CCL2 overexpression led to a selective increase in LIF mRNA and neuronal phosphorylated STAT3 (pSTAT3) in L5 DRGs, with no change in expression seen in other RAGs such as GAP-43. Blockade of STAT3 phosphorylation by each of two different inhibitors prevented the increase in neurite outgrowth. Thus, CCL2 overexpression is sufficient to induce macrophage accumulation in uninjured L5 DRGs and increase the regenerative capacity of DRG neurons via a STAT3-dependent mechanism.
神经炎症在轴突切断术后外周神经的再生中起关键作用。坐骨神经损伤会导致L5背根神经节中巨噬细胞大量聚集,而背根损伤时则不会出现这种现象。我们最近证明,轴突切断的细胞体周围的这种聚集是外周预处理损伤反应发生所必需的。在这里,我们探讨了在未受伤小鼠的背根神经节神经元中特异性过表达单核细胞趋化因子CCL2是否足以引起巨噬细胞聚集并促进再生,或者是否需要其他损伤衍生信号。通过鞘内注射AAV5-EF1α-CCL2,这种注射导致CCL2 mRNA表达和L5背根神经节中巨噬细胞聚集呈时间依赖性增加,在注射后3周达到最大反应。这些变化导致背根神经节外植体和分离细胞培养物中神经突生长出现类似预处理的增加。这种再生增加依赖于CCL2通过其主要受体CCR2发挥作用。当在CCR2基因敲除小鼠中过表达CCL2时,未观察到巨噬细胞聚集和再生增强。为了探讨CCL2过表达增强再生的机制,我们检测了这些动物中再生相关基因的表达是否升高。令人惊讶的是,我们发现CCL2过表达导致L5背根神经节中白血病抑制因子(LIF)mRNA和神经元磷酸化信号转导子和转录激活子3(pSTAT3)选择性增加,而其他再生相关基因如生长相关蛋白43(GAP-43)的表达没有变化。两种不同抑制剂对STAT3磷酸化的阻断均阻止了神经突生长的增加。因此,CCL2过表达足以在未受伤的L5背根神经节中诱导巨噬细胞聚集,并通过STAT3依赖性机制增加背根神经节神经元的再生能力。