CCR7的表达及与配体CCL19对鼠小胶质细胞生物学功能的影响
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摘要
趋化因子与表达于细胞表面的受体共同作用发挥其生物学功能。趋化因子受体均为G蛋白偶联受体，与趋化因子结合后，通过G蛋白变构而使受体磷酸化，进行信号转导。CCR7是由EBV诱导后发现的基因，被认为是EBV在B淋巴细胞上作用的调节因子。CCR7受体表达于各种淋巴组织，并激活T、B淋巴细胞。作为免疫系统中一个重要的归巢受体，CCR7不仅控制B细胞和T细胞归巢、DCs穿过高内皮静脉，而且还能把它们正确定位在外周淋巴器官的T、B细胞区。CCR7可能对趋化成熟DCs进入淋巴管非常重要。SLC是由淋巴管内皮细胞产生的，从淋巴管，DCs回流到本地LN，最终定位于T细胞区。这一最后定位过程可能被ELC控制，因为ELC被定居在此的成熟DCs产生。在免疫刺激或炎症刺激后，DCs从外周组织迁移到淋巴器官，提呈抗原。在此过程中由非成熟DCs转变为成熟DCs，在成熟过程中下调两个CC趋化因子受体：CCR1、CCR5，对其配体(HCC-1，MIP-1β)的趋化活性消失，但上调CCR7表达。对呈现在炎症位点或免疫位点的趋化因子的反应的抑制使其离开外周组织，对SLC、ELC的反应引导其定位到淋巴器官。由于CCR7重要性，研究者在不同的细胞对这一趋化因子受体进行大量研究，然而到目前为止，还没有关于小胶质细胞中CCR7表达情况的研究。近年来发现，CCL21与其中一个CXCR3互作诱导小胶质细胞的趋化。进一步研究发现，在CNS中这一信号系统不可能发挥重要作用。
本试验用3d内gcd-10野生型小鼠作为实验材料，首次采用RT-PCR和Western blotting方法，发现活化的小胶质细胞表达CCR7，然而在静息状态下并不表达。LPS活化的原代小胶质细胞CCR7的表达量随时间和LPS处理剂量的变化而变化。同时，我们还发现Cell Debris处理的小胶质细胞中也有CCR7的表达，而IFN-γ激活的小胶质细胞并不表达。
TGF-β1与多种细胞因子的产生有关：TGF-β1能抑制各种细胞因子的表达。本试验使用TGF-β1处理LPS活化的小胶质细胞发现，TGF-β1可明显抑制CCR7 mRNA的表达。本试验还使用了PI3K、MEK／ERK和p38信号通路的阻断剂LY294002、U0126和SB203580处理原代培养的小胶质细胞，结果没有检测到LY294002或SB203580影响CCR7 mRNA的表达，U0126上调CCR7的表达。
本试验还检测了LPS处理的小胶质细胞TNF-α和iNOS mRNA表达。LPS活化的小胶质细胞表达前炎症因子TNF-α和iNOS mRNA，且TNF-αmRNA的表达早于iNOS mRNA。LPS处理6h，TNF-αmRNA在所选取的处理时间表达量最高，24h的表达量与处理前的表达水平相当；iNOS的表达峰值出现在LPS处理6h，并且直到24h iNOS mRNA仍然维持高水平表达
活化的小胶质细胞表达CCR7促使我们研究该受体与其配体CCL19对细胞生物学功能的影响。结果表明，LPS处理的小胶质细胞的死亡率约为20％，当加入CCL19后，细胞死亡率降低到10％左右，提示CCL19能减少约50％LPS处理的小胶质细胞死亡率，因此CCL19与其受体作用可以降低细胞的死亡率。我们还发现，在LPS处理的小胶质细胞中加入CCL19后，可显著提高细胞的吞噬作用和胞内Ca~(2+)浓度。
结论：LPS或Cell Debris活化的小胶质细胞表达CCR7，而处于静息状态的小胶质细胞并不表达该趋化因子受体；CCR7的配体MIP-3β可提高LPS处理的小胶质细胞的存活率、吞噬作用以及胞内Ca~(2+)浓度。
Chemokines deliver their activity by interacting with cell surface expressed chemokine receptors that belong to the family of seven transmembrane domain G protein-coupled rhodopsin-like receptors. CCR7 was identified as a gene induced by the Epstein-Barr virus (EBV), and is thought to be a mediator of EBV effects on B lymphocytes. Previous results told us that such receptor is expressed in various lymphoid tissues and activates B and T lymphocytes. Important functions of CCR7 have been shown to control the migration of memory T cells to inflamed tissues and stimulate dendritic cell maturation. Because of the importance of such receptor, many researchers have investigated CCR7 in many kinds of cells. However, no report has been made regarding the presence of CCR7, the receptor for CCL19 and CCL21 in microglia; instead, CCL21 is found to induce chemotactic behavior by interacting with CXCR3 in microglia. Moreover, recent studies have shown that it is highly unlikely that this signaling system plays a role in inflammation in the CNS.
Experimental animals used in this study were gcd-10 wild type mouse. Our present results provide the first evidence that RT-PCR and Western blotting analysis showed CCR7 expression was detected in activated primary cultures of microglia, but not in the resting mocroglia. Primary cultures of microglia stimulated with LPS express CCR7 mRNA in time and dose-dependent way. CCR7 mRNA expression was also found in cultured microglia treated by cell debris. LPS, but not IFN-γinduced CCR7 expression in microglial cell. Similar to previous results, expression of CXCR3 mRNA was found in the cultured microglial cells.
TGF-βM has known to be associated with production of various kinds of cytokines. TGF-β1 inhibits various kinds of cytokine mRNA expression. In present study, we found that LPS-induced expression CCR7 mRNA is almost suppressed by TGF-β1. However, LY294002, a PI3K inhibitor, has no effect on LPS-induced CCR7 mRNA expression in cultured microglia, suggesting that TGF-β1 shows its activity not through PI3K signal pathway. Previously, it has been reported that induction of CCR7 expression in thymocytes and RLm6 depends on activation of the MEK-ERK signaling pathway and an increase in intracellular Ca~(2+) levels. Here we examined the effect of U0126, an inhibitor of the extracellular signal-regulated kinase kinase (ERKK/MEK), on expression of CCR7 mRNA in activated microglia. Interestingly, however, CCR7 mRNA expression was up-regulated after adding the U0126. In the present study we also investigated other pathways such as p38, which is possibly responsible for the induction of CCR7 mRNA expression in the activated primary cultures of microglia. Our data indicated that CCR7 mRNA expression does not due to the activation of the ERKK/MEK or p38. Although the induction mechanism of CCR7 expression in activated microglia requires further investigation, the cell type, experimental condition-dependent effect and some other factors, such as NF-κB, the interplay of NF-κB with MAPKs or the interplay within the MAPKs, could be due to mechanism for the induction of CCR7 mRNA expression.
Similar to previous results, TNF-αand iNOS mRNA expression were detected in LPS stimulated microglia.
The findings that CCR7 mRNA is expressed in activated microglia led us to examine effects of its ligands. In the present study, we examined the effects of CCL19 on LPS-treated primary microglial cells culture. Percent of Cell death was almost 20% in LPS-treated cells. This percentage was consistently reduced to about 10% when the cells were cultured in the presence of 0.lμg/ml CCL19, implying that CCL19 reduced about 50% of dead cells. Our results clearly show that CCL19 is able to augment the function of phagocytosis for LPS-stimulated microglia. After adding the CCL19 in LPS-stimulated microglia, phagocytosis was augmented, indicating that the interaction between CCR7 and its ligand, CCL19 enhances phagocytosis of microglia. With the adding of CCL19 a transient increase in intracellular calcium was detected in the cultured microglia under activated condition but not under normal condition, indicating that level of Ca~(2+) influx can be elevated by CCL19 in activated microglia.
In conclusion, CCR7 is barely expressed in microglia under normal conditions, but markedly up-regulated upon their activation. Indeed, its ligand CCL19 augmented phagocytosis in activated microglia, suggesting that under activated conditions the CCL19/CCR7 interaction becomes predominant, which may play a role in regulating chemokine receptors and cytokine production, cell survival, phagocytosis and transient increase in intracellular calcium.
本试验用3d内gcd-10野生型小鼠作为实验材料，首次采用RT-PCR和Western blotting方法，发现活化的小胶质细胞表达CCR7，然而在静息状态下并不表达。LPS活化的原代小胶质细胞CCR7的表达量随时间和LPS处理剂量的变化而变化。同时，我们还发现Cell Debris处理的小胶质细胞中也有CCR7的表达，而IFN-γ激活的小胶质细胞并不表达。
TGF-β1与多种细胞因子的产生有关：TGF-β1能抑制各种细胞因子的表达。本试验使用TGF-β1处理LPS活化的小胶质细胞发现，TGF-β1可明显抑制CCR7 mRNA的表达。本试验还使用了PI3K、MEK／ERK和p38信号通路的阻断剂LY294002、U0126和SB203580处理原代培养的小胶质细胞，结果没有检测到LY294002或SB203580影响CCR7 mRNA的表达，U0126上调CCR7的表达。
本试验还检测了LPS处理的小胶质细胞TNF-α和iNOS mRNA表达。LPS活化的小胶质细胞表达前炎症因子TNF-α和iNOS mRNA，且TNF-αmRNA的表达早于iNOS mRNA。LPS处理6h，TNF-αmRNA在所选取的处理时间表达量最高，24h的表达量与处理前的表达水平相当；iNOS的表达峰值出现在LPS处理6h，并且直到24h iNOS mRNA仍然维持高水平表达
活化的小胶质细胞表达CCR7促使我们研究该受体与其配体CCL19对细胞生物学功能的影响。结果表明，LPS处理的小胶质细胞的死亡率约为20％，当加入CCL19后，细胞死亡率降低到10％左右，提示CCL19能减少约50％LPS处理的小胶质细胞死亡率，因此CCL19与其受体作用可以降低细胞的死亡率。我们还发现，在LPS处理的小胶质细胞中加入CCL19后，可显著提高细胞的吞噬作用和胞内Ca~(2+)浓度。
结论：LPS或Cell Debris活化的小胶质细胞表达CCR7，而处于静息状态的小胶质细胞并不表达该趋化因子受体；CCR7的配体MIP-3β可提高LPS处理的小胶质细胞的存活率、吞噬作用以及胞内Ca~(2+)浓度。
Chemokines deliver their activity by interacting with cell surface expressed chemokine receptors that belong to the family of seven transmembrane domain G protein-coupled rhodopsin-like receptors. CCR7 was identified as a gene induced by the Epstein-Barr virus (EBV), and is thought to be a mediator of EBV effects on B lymphocytes. Previous results told us that such receptor is expressed in various lymphoid tissues and activates B and T lymphocytes. Important functions of CCR7 have been shown to control the migration of memory T cells to inflamed tissues and stimulate dendritic cell maturation. Because of the importance of such receptor, many researchers have investigated CCR7 in many kinds of cells. However, no report has been made regarding the presence of CCR7, the receptor for CCL19 and CCL21 in microglia; instead, CCL21 is found to induce chemotactic behavior by interacting with CXCR3 in microglia. Moreover, recent studies have shown that it is highly unlikely that this signaling system plays a role in inflammation in the CNS.
Experimental animals used in this study were gcd-10 wild type mouse. Our present results provide the first evidence that RT-PCR and Western blotting analysis showed CCR7 expression was detected in activated primary cultures of microglia, but not in the resting mocroglia. Primary cultures of microglia stimulated with LPS express CCR7 mRNA in time and dose-dependent way. CCR7 mRNA expression was also found in cultured microglia treated by cell debris. LPS, but not IFN-γinduced CCR7 expression in microglial cell. Similar to previous results, expression of CXCR3 mRNA was found in the cultured microglial cells.
TGF-βM has known to be associated with production of various kinds of cytokines. TGF-β1 inhibits various kinds of cytokine mRNA expression. In present study, we found that LPS-induced expression CCR7 mRNA is almost suppressed by TGF-β1. However, LY294002, a PI3K inhibitor, has no effect on LPS-induced CCR7 mRNA expression in cultured microglia, suggesting that TGF-β1 shows its activity not through PI3K signal pathway. Previously, it has been reported that induction of CCR7 expression in thymocytes and RLm6 depends on activation of the MEK-ERK signaling pathway and an increase in intracellular Ca~(2+) levels. Here we examined the effect of U0126, an inhibitor of the extracellular signal-regulated kinase kinase (ERKK/MEK), on expression of CCR7 mRNA in activated microglia. Interestingly, however, CCR7 mRNA expression was up-regulated after adding the U0126. In the present study we also investigated other pathways such as p38, which is possibly responsible for the induction of CCR7 mRNA expression in the activated primary cultures of microglia. Our data indicated that CCR7 mRNA expression does not due to the activation of the ERKK/MEK or p38. Although the induction mechanism of CCR7 expression in activated microglia requires further investigation, the cell type, experimental condition-dependent effect and some other factors, such as NF-κB, the interplay of NF-κB with MAPKs or the interplay within the MAPKs, could be due to mechanism for the induction of CCR7 mRNA expression.
Similar to previous results, TNF-αand iNOS mRNA expression were detected in LPS stimulated microglia.
The findings that CCR7 mRNA is expressed in activated microglia led us to examine effects of its ligands. In the present study, we examined the effects of CCL19 on LPS-treated primary microglial cells culture. Percent of Cell death was almost 20% in LPS-treated cells. This percentage was consistently reduced to about 10% when the cells were cultured in the presence of 0.lμg/ml CCL19, implying that CCL19 reduced about 50% of dead cells. Our results clearly show that CCL19 is able to augment the function of phagocytosis for LPS-stimulated microglia. After adding the CCL19 in LPS-stimulated microglia, phagocytosis was augmented, indicating that the interaction between CCR7 and its ligand, CCL19 enhances phagocytosis of microglia. With the adding of CCL19 a transient increase in intracellular calcium was detected in the cultured microglia under activated condition but not under normal condition, indicating that level of Ca~(2+) influx can be elevated by CCL19 in activated microglia.
In conclusion, CCR7 is barely expressed in microglia under normal conditions, but markedly up-regulated upon their activation. Indeed, its ligand CCL19 augmented phagocytosis in activated microglia, suggesting that under activated conditions the CCL19/CCR7 interaction becomes predominant, which may play a role in regulating chemokine receptors and cytokine production, cell survival, phagocytosis and transient increase in intracellular calcium.
引文
李怡，何成．胶质细胞介导性激素的神经系统作用．生理科学进展．2002，33：35-38．
万选才，杨天祝，徐承熹主编．现代神经生物学．北京医科大学中国协和医科大学联合出版社1999，91-96．
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