神經元細胞接受刺激或受到損傷後會釋放大量的麩胺酸而造成興奮性神經毒性反應。中樞神經系統內含量最多的膠質細胞為星狀膠細胞，可以透過膜上特定表現的麩胺酸轉運蛋白如麩胺酸/天門冬胺酸轉運蛋白(glutamate/
aspartate transporter, GLAST /EAAT1)與麩胺酸轉運蛋白-1
(glutamate transporter-1, GLT-1/EAAT2)來回收胞外高濃度麩胺酸，藉此維持突觸間麩胺酸濃度的恆定。其中以GLT-1最為重要，前腦區域約90%胞外麩胺酸都是由此所回收，然而在大腦發育過程中卻不易偵測到GLT-1的表現。視黃酸(retinoic acid，RA)在脊椎動物神經系統發育與維持扮演重要角色。研究指出，視黃酸會調控成體間葉幹細胞(adult mesenchymal stem cells)的GLT-1基因表現。本實驗室先前以軟體(Footer)預測發現，視黃酸受體(RXR)與GLT-1啟動子可能有七個結合位。基於此，本篇的研究目的主要是想要探討，視黃酸是否也參與調控星狀膠細胞的GLT-1表現。結果發現，視黃酸會抑制星狀膠細胞的GLT-1 mRNA與蛋白質表現。EMSA結果指出，視黃酸可能會經由GLT-1啟動子的RARE 6片段調控GLT-1基因表現，猜測視黃酸可能是透過轉錄調控抑制星狀膠細胞的GLT-1表現。文獻指出，雙丁醯環磷腺苷(dbcAMP)會提升GLT-1的表現量並促進其運送至星狀膠細胞膜上，因此接著探討視黃酸對於dbcAMP處理星狀膠細胞的GLT-1表現影響。在F-actin染色實驗觀察到，視黃酸會使dbcAMP處理細胞變得比較扁平。而視黃酸也會抑制dbcAMP處理細胞的GLT-1 mRNA表現並進一步抑制其麩胺酸回收能力。在慢病毒基因傳輸系統發現，視黃酸降低dbcAMP處理星狀膠細胞觸支上的GLT-1表現。另外我們也觀察到視黃酸會提升dbcAMP處理細胞的PKC活性，因此我們使用兩種PKC抑制劑Bis I與rottlerin來確認視黃酸是否會透過PKC的活化來抑制GLT-1表現。結果指出，處理PKC抑制劑後，視黃酸就無法降低dbcAMP處理細胞的GLT-1膜蛋白表現。另外我們也發現視黃酸影響星狀膠細胞的GLT-1膜蛋白分布可能也與dynamin與clathrin的調控有關。綜合以上結果指出，視黃酸可能會透過RXR所誘導的轉錄調控抑制GLT-1表現，或者是藉由PKC的活化造成GLT-1膜蛋白表現下降，而這樣的現象可能與dynamin與clathrin的參與有關。

In the mammalian central nervous system (CNS), astrocytes are responsible for clearance of excess extracellular glutamate molecules through glutamate transporter-1 (GLT-1)/EAAT-2 and Na+-dependent glutamate/aspartate
transporter (GLAST). GLT-1 is the most important glutamate transporter subtype sice it is responsible for 90% of forebrain glutamate uptake in adult, wherase GLT-1 expression is undetected in developing brain. Retinoic acid (RA), a potent regulator of differentiation in the nervous system, mediates GLT-1 expression in the adult mesenchymal stem cells. Rat GLT-1 promoter sequence contains several retinoid X receptor (RXR) DNA binding regions. Accordingly, we attempted to examine whether RA signaling is involved in the regulation of GLT-1 expression in primary rat astrocytes. Our results showed that treatment with RA for 24 h significantly reduced GLT-1 mRNA and GLT-1 membrane protein levels in astrocytes. Electrophoresis motility shift assay (EMSA) indicated that the RXR-DNA binding activity was increased in astrocytes after treatment with RA for 1 h, suggesting that the decline of GLT-1 levels in RA-treated astrocytes was mediated at the transcriptional levels. Given that intracellular cAMP is involved in GLT-1 expression and cellular trafficking in astrocytes, the effect of RA on dbcAMP-treated astrocytes was further examined. F-actin staining indicated that RA caused dbcAMP-treated astrocytes to become more flat. Similar to the condition in the absence of dbcAMP, RA significantly inhibited GLT-1 mRNA expression in dbcAMP-treated astrocytes and suppressed their glutamate uptake activity. Through lentiviral gene delivery approach, GLT-1levels in the processes of dbcAMP-treated astrocytes were attenuated by RA. Based on the finding that RA induced protein kinase C (PKC) activation in dbcAMP-treated astrocytes, the two inhibitors of PKC, Bis I and rottlerin, were used to determine the involvement of PKC in RA-induced GLT-1 reduction. Indeed, pretreatment with the PKC inhibitors, RA failed to reduce GLT-1 membrane protein levels in dbcAMP-treated astrocytes. Furthermore, we found that dynamin and clatherin were invovled in RA-induced redistribution of GLT-1 in astrocytic cell surface. In summary, we conclude that RA reduces GLT-1 levels in astrocytes not only through RXR-mediated inhibition at its transcriptional level, but also triggers the activation of PKC to reduce cell surface levels of GLT-1 possibly via dynamin and clathrin-dependent pathways.

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