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研究生:金亭佑
研究生(外文):Ting-Yu Chin
論文名稱:(1)豬大動脈平滑肌細胞胞內鈣訊號之研究(2)類鴉片藥物脫敏作用之研究
論文名稱(外文):(1) Ca2+ signaling induced by ATP and sphingosylphosphorylcholinein porcine aortic smooth muscle cells (2) Studies of desensitization mechanism of opioid drugs
指導教授:闕小輝
指導教授(外文):Sheau-Huei Chueh
學位類別:博士
校院名稱:國防醫學院
系所名稱:生命科學研究所
學門:生命科學學門
學類:生物學類
論文種類:學術論文
論文出版年:2002
畢業學年度:90
語文別:中文
中文關鍵詞:鈣離子類鴉片受體
外文關鍵詞:calciumopioid receptor
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  • 收藏至我的研究室書目清單書目收藏:1
(1)
在豬大動脈平滑肌細胞中,利用鈣偵測劑 fura-2,測定經 ATP或sphingosyl- phosphorylcholine (SPC) 重複刺激所引起的胞內鈣離子濃度增加。
ATP 是經由鈣庫釋放鈣以及胞外鈣內流造成胞內鈣離子濃度增加。前者是磷酸脂酶 C (phospholipase C) 活化作用所引發的;後者明顯是經由受體本身即是鈣離子通道 (receptor-operated Ca2+ channel; ROC),而非鈣庫調控的鈣離子通道(store-operated Ca2+ channel; SOC) 或電感性鈣離子通道 (voltage-operated Ca2+ channel; VOC。更進一步實驗顯示,P2X5 受體負責ATP所誘發的胞外鈣內流。
ATP 重複刺激可產生一具有再現性的胞內鈣離子濃度增加,但此現象會因胞外鈣離子移除,或使用U-73122 或 thapsigargin 抑制胞內鈣庫釋放鈣而受阻。前者若將胞外鈣離子再加回,則再現性又恢復。
SPC只能造成鈣釋放,所以重複SPC刺激所引起的胞內鈣離子增加的大小會逐漸減小。然而當蛋白質激酶 C (protein kinase C) 被抑制時,其可增加SPC誘發胞外鈣內流,因此重複 SPC 刺激也產生具有再現性的胞內鈣離子濃度增加。
總而言之,當胞外鈣內流受阻或將胞內鈣釋放抑制,皆會使ATP重複刺激所引發的鈣上升逐漸減弱。當在正常生理狀況下,每一次ATP刺激的整個胞內鈣離子濃度增加量是維持相同的,這說明當受刺激時,鈣庫會經由胞外鈣內流再被補充。而 SPC完全是經由鈣釋放引起胞內鈣離子濃度增加,且其反應會隨著重複刺激而逐漸減小,但此下降反應會因活化胞外鈣內流而被避免。這更進一步支持經胞外鈣內流回補胞內鈣庫對持續鈣訊息途徑的重要性。
(2)
為了瞭解磷酸化作用在δ-類鴉片受體脫敏現象中所扮演的角色,我們利用定點突變方式,將δ-類鴉片受體位在胞質中的 C 端五個可能磷酸化的殘基進行突變,得到四種突變組:分別為 T335A,S344A,T358A,T361A+S363A。再使用 pcDNA3.1 表現載體,將野生型和突變型δ-類鴉片受體送入 HEK293 細胞株中表現。無論野生型或突變型δ-類鴉片受體組,其給予致活劑五分鐘對於 forskolin 所增加的 cAMP 生成量之抑制作用類似,約 50-70%。然而,在野生型或T335A突變型組,如果將細胞先曝露在δ-類鴉片受體的致活劑四小時後,再測其抑制cAMP 生成量的作用,其抑制能力減弱,只剩下將近18-25%。但S344A,T358A,T361A+S363A 這三組在慢性致活劑處理下,並沒有明顯的脫敏現象,其在致活劑處理四小時後,仍能維持將近 60% 的抑制效果,與其急性致活劑處理時的抑制情形類似。
另一方面前處理 mitogen-activated protein kinase kinase (MEK) 的抑制劑 PD098059,在野生型或T335A突變型組,會使慢性致活劑處理造成的cAMP 生成量之抑制作用的脫敏現象消失。而且受體連結分析和免疫螢光共軛焦顯微鏡觀察的證據顯示,δ-類鴉片受體 internalization 需要 MAPK 活性。
除了抑制cAMP 生成作用,δ-類鴉片受體活化時也會引起胞內鈣離子濃度增加,其誘發鈣離子上升是短暫的,且在二分鐘內即回到基礎。無論野生型或所有突變型δ-類鴉片受體組,給予致活劑刺激引起的鈣上升皆類似,且都是將近 5-15% 的細胞在給予致活劑 DADLE 會造成 10-20 秒的鈣離子增加。前處理 PD098059,會增加具有反應的細胞數百分比,由5-15% 增至32-46%。此現象在野生型或所有突變型δ-類鴉片受體組均是類似的。
這些結果指出,在δ-類鴉片受體訊息途徑的調控機轉是多重的。Gi 蛋白連接的抑制 cAMP訊息途徑之脫敏現象,是經由 MAPK。δ-類鴉片受體 C端 S/T 位置的磷酸化也參與。而δ-類鴉片受體活化後所激活的MAPK會再回饋抑制磷酸脂酶C 的訊息途徑,但此途徑之調控機轉中並不需要δ-類鴉片受體 C 端磷酸化。
我們也想了解長期施打嗎啡的懷孕母鼠,其子代的海馬迴神經細胞中之 NMDA 受體活性是否會改變,所以以出生一天之白鼠為材料。
利用鈣螢光偵測劑fura-2,我們發現在海馬迴神經細胞中,在有 10μM 甘胺酸 (glycine) 和沒有胞外鎂離子的情況下,麩胺酸 (glutamate) 和 N-methyl-D-aspartate (NMDA) 會誘發胞內鈣離子濃度增加,且此增加是劑量依賴性的。在控制組方面,麩胺酸和NMDA的EC50 分別為1.2μM 和6.6μM。S-nitrosoglutathione (GSNO) 是一種一氧化氮提供者,同樣也會引起胞內鈣離子濃度增加;其大小相當於 NMDA 所引發的鈣離子變化,且可受到鎂離子抑制。然而其他一氧化氮提供者,例如 3-morpholinosydnonimine (SIN-1), S-nitroso-N-acetylpenicillamine (SNAP), and sodium nitroprusside (SNP) 卻無法使胞內鈣離子濃度上升。由此可知,GSNO可能是經由 GSH 活化 NMDA 受體的。爾後的確發現 GSH 可促使胞內鈣離子濃度增加和麩胺酸釋放,其EC50 分別為490μM 和680μM。GSH 引發胞內鈣離子濃度上升的能力會因 NMDA 受體的拮抗劑 AP-5或鎂離子的加入而被抑制。但其他 GSH 衍生物如S-methylGSH、 S-ethylGSH、 S-propylGSH、 S-butyl GSH、 Glu-Cys 、 Cys-Gly,或與GSH 同樣具有還原能力的 dithiothreitol (DTT) and mercaptoethanol 皆無法使胞內鈣離子濃度增加。而細胞處理N-ethylmaleimide (NEM) 後,也不會改變 NMDA 及 GSNO 引發鈣離子濃度上升的作用,所以排除 sulfhydryl group 參與 NMDA 受體通道開啟的可能性。綜合以上結果可知,GSH 可能是 NMDA 受體的致活劑。
綜合所有實驗結果可知,有或無嗎啡處理對GSH 促使胞內鈣離子濃度增加和麩胺酸釋放的能力上,並沒有顯著差異。因此,長期施打嗎啡的懷孕母鼠,對於其子代的海馬迴神經細胞中,GSH 經 NMDA 受體所引發的胞內鈣離子濃度上升並沒有明顯的影響。

(1)
The increase in the cytosolic Ca2+ concentration ([Ca2+]i) following repetitive stimulation with ATP or sphingosylphosphorylcholine (SPC) in single smooth muscle cells was investigated using the Ca2+ indicator, fura-2.
The ATP-induced Ca2+ increase resulted from both Ca2+ release and Ca2+ influx. The former was stimulated by phospholipase C activation, while the latter occurred predominantly via the receptor-operated Ca2+ channels (ROC), rather than the store-operated Ca2+ channels (SOC) or the voltage-operated Ca2+ channels (VOC). Furthermore, the P2X5 receptor was shown to be responsible for the ATP-induced Ca2+ influx.
A reproducible [Ca2+]i increase was induced by repetitive ATP stimulation, but was abolished by removal of extracellular Ca2+ or inhibition of intracellular Ca2+ release using U-73122 or thapsigargin, and was restored by Ca2+ readdition in the former case.
SPC only caused Ca2+ release, and the amplitude of repetitive SPC- induced [Ca2+]i increases declined gradually. However, a reproducible [Ca2+]i increase was seen in cells in which protein kinase C being inhibited, which increased the SPC-induced Ca2+ influx, rather than IP3 generation.
In conclusion, although the amplitude of the ATP-increase Ca2+ release, measured when Ca2+ influx was blocked, or of the Ca2+ influx when Ca2+ release was blocked, progressively decreased following repetitive stimulation, the overall [Ca2+]i increase for each stimulation under physiological conditions remained the same, suggesting that the Ca2+ stores were replenished by an influx of Ca2+ during stimulation. The SPC-induced [Ca2+]i increase resulted solely from Ca2+ release and decrease gradually following repetitive stimulation, but the decrease could be prevented by stimulating Ca2+ influx, further supporting involvement of the intracellular Ca2+ stores in Ca2+ signalling.
(2)
To characterize the role of phosphorylation in the process of desensitization of delta opioid receptor (DOR), using site-directed mutagenesis, five potential phosphorylation residues in the C-terminal cytosolic tail of DOR were mutated and four mutants were obtained, T335A, S344A, T358A and T361A + S363A. Using expression vector pcDNA3.1, wild type and mutants of DOR were transfected and expressed in HEK293 cells. The inhibitory effect of DOR on cAMP generation was still seen all groups have been exposed to agonist for 5 min, by approximately 50-70%. However, it was reversed after exposure cells with agonists for 4 h; the inhibition was only approximately 18 — 22% in wild type and T335A mutant. Whereas, in S344A, T358A and T361A + S363A mutants, desensitization of inhibition of cAMP generation after chronic agonist treatment were not seen contrasting with wild type; agonist-induced inhibition of cAMP generation remained approximately 60% after prior incubation with agonist for 4 h, similar to that of acute effect.
On the other hand, pretreatment of cells with PD098059, an inhibitor of mitogen-activated protein kinase kinase (MEK), in wild type or T335A mutant, which blocks the desensitization of inhibition of cAMP generation after chronic agonist treatment. Both receptor binding assay and immunofluorescence confocal microscopic evidence suggests that receptor internalization require MAPK activity.
In addition to inhibition of cAMP generation, activation of DOR also induced an increase in cytosolic Ca2+ concentration ([Ca2+]i). DOR-induced [Ca2+]i increase was transient and [Ca2+]i returned to the basal level within 2 min after stimulation. All mutants displayed similar characteristics as wild type in [Ca2+]i increase the amplitude of increased [Ca2+]i and the percentage of responding cells upon stimulation. Approximately 5-15% of cells regardless of the wild type or mutants responded to DADLE in which [Ca2+]i increased 10 —20 sec after DADLE addition. Pretreatment of cells with PD098059 increased the percentage of responding cells upon stimulation; DADLE-induced [Ca2+]i increase was observed in approximately 32-46% of cells. The enhancement effect of PD098059 in opioids-induced Ca2+ signaling was identical in both wild type and mutants cells.
These data indicate that there exists distinct regulatory mechanisms in DOR signaling pathways. Desensitization of Gi-coupled cAMP signaling pathway is governed by both MAPK activity and the phosphorylation of C-terminal S/T sites of DOR. Activation of DOR stimulates MAPK which in turn may feedback inhibits the phospholipase C signaling pathway and is independent to the C-terminal phosphorylation of DOR.
We determined whether continuous exposure to morphine beginning from prenatal period will alter the activity of NMDA receptor in the hippocampal neuron of rat offspring.
Using fluorescent Ca2+ indicator fura-2, we found that in cultured hippocampal neurons, in the presence of 10μM glycine and absence of Mg2+ in the bathing buffer, glutamate or N-methyl-D-aspartate (NMDA) dose-dependently induced an increase of the cytosolic Ca2+ concentration ([Ca2+ ]i). In control group, the EC50 value for glutamate and NMDA were 1.2μM and 6.6μM, respectively. S-nitrosoglutathione (GSNO), a nitric oxide (NO) donor, also evoked a [Ca2+]i increase; the amplitude of it was identical to that of NMDA and inhibited by Mg2+, while other NO donors, including 3-morpholinosydnonimine (SIN-1), S-nitroso-N-acetylpenicillamine (SNAP), and sodium nitroprusside (SNP), could not suggesting that glutathione (GSH) may activate NMDA receptor. Indeed, GSH induced [Ca2+]i increase and glutamate release with EC50 value of 490μM and 680μM, respectively, and GSH-induced [Ca2+ ]i increase is inhibited by AP-5 or Mg2+ addition, while other GSH derivatives, including S-methylGSH, S-ethylGSH, S-propylGSH, S-butyl GSH, Glu-Cys and Cys-Gly, dithiothreitol (DTT) and mercaptoethanol, had no effect. Treatment of cells with N-ethylmaleimide (NEM) did not alter the action of NMDA and GSNO on increase precluding the involvement of sulfhydryl group in the channel opening. In conclusion, GSH may act as an agonist for NMDA receptor.
There is no significant difference in GSH-induced [Ca2+]i increase and glutamate release between control group and morphine group. Thus, chronic maternal morphine exposure does not modulate GSH -induced [Ca2+]i increase via NMDA receptor in the hippocampal neurons of the offsprings.

目錄
頁次
目錄 ............................. .I
圖、表目錄............................ .II
縮寫表.............................. .V
第一部分、豬大動脈平滑肌細胞胞內鈣訊號之研究........... .1
中文摘要............................ .2
英文摘要............................ .3
第一章緒論............................ .4
第二章實驗材料與方法....................... .15
第三章結果........................... .19
第四章討論............................ .35
參考文獻............................ .40
第二部分、類鴉片藥物脫敏作用之研究................ .54
中文摘要............................ .55
英文摘要............................ .57
第一章緒論............................ .60
第二章實驗材料與方法....................... .68
第三章結果........................... .80
第四章討論........................... .109
參考文獻............................ .116
附錄 ............................. .126

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