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研究生:蔡鳳如
研究生(外文):Feng-Ru Tsai
論文名稱:IGF-1促進胚胎發育時期運動神經肌肉細胞突觸之自發性神經傳導物質釋放之研究
論文名稱(外文):potentiation of spontaneous transmitter release by IGF-1 at developing neuromuscular synapse.
指導教授:劉昭成
指導教授(外文):Jau-Cheng Liou
學位類別:碩士
校院名稱:國立中山大學
系所名稱:生物科學系研究所
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
論文頁數:59
中文關鍵詞:神經與肌肉細胞突觸訊息傳遞路徑
外文關鍵詞:synapseXenopusIGF-1
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  • 被引用被引用:2
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 神經細胞與肌細胞形成突觸的過程非常複雜,這過程中除了細胞彼此間藉著細胞黏著因子的直接接觸產生交互作用之外,許多訊號分子如ATP、NO、HPETE的出現及一些神經滋養因子的產生會進一步穩定整個突觸結構。在本論文中我們將探討Insulin-like growth factor-I (IGF-I) 在胚胎時期對神經細胞與肌細胞突觸形成過程所扮演的角色。
IGFs是在1970年代被cloned出來,隨著眾多實驗的進行發現IGFs除了在生理代謝上有著類似胰島素的功能,在週圍及中樞神經系統也扮演著生長、分化及再生的重要角色,由於目前對於IGF-I在胚胎早期神經細胞與肌細胞間突觸形成過程中所扮演的角色所之甚少,因此本實驗利用非洲爪蟾的神經細胞與肌細胞的混合培養( Xenopus nerve-muscle co-culture ),以whole-cell patch clamp的電生理記錄的方式來探討IGF-I在胚胎發育早期突觸形成過程中所扮演的角色以及這中間可能的訊息傳遞路徑。藉由在肌細胞記錄神經所釋放的ACh之後,ACh打開肌細胞上ACh receptor而造成自發性電流,我們可以清楚的觀察到神經細胞的活性,當我們在培養皿中加入IGF-I之後約經15分鐘後我們發現自發性神經傳導物質釋放的頻率有顯著的增加。由於神經細胞釋放神經傳導物質和神經末梢內鈣離子的濃度有很大的關係,因此我們設計一系列實驗來釐清鈣離子的來源。實驗結果顯示在Ca2+ free Ringer以及鈣離子通道阻斷劑Cd2+的存在下IGF-I對神經活性的促進作用依然存在,顯示IGF-I對神經活性的促進作用所需的鈣離子來源不是來自細胞外,而是由細胞內的鈣離子儲存池所提供。此外,為了更進一步證實IGF-I的作用和鈣離子儲存池的關係,我們利用鈣離子儲存池上的鈣離子通道阻斷劑:IP3 receptor inhibitor (XeC, 2-APB) 及ryanodine receptor inhibitor (TMB-8) 或是用鈣離子儲存池的排空劑thapsigargin以阻斷細胞內鈣離子的來源,實驗結果一致顯示在沒有細胞內的鈣離子來源的情況下IGF-I便無法再促進神經活性。至於IGF-I的那些訊息傳遞路徑和促進神經活性有關?目前已知IGF-I有三條訊息傳遞路徑─PI 3-kinase、PLCg和MAP kinase。這次的實驗結果發現,IGF-I的作用會因PI 3-kinase和PLCg的活性被抑制而消失,但對於MAP kinase的抑制作用沒有明顯的影響。
當神經末梢內鈣離子濃度升高時,Ca2+會透過和calmoldulin形成複合物而將Ca2+/calmoldulin-dependent protein kinase (CaMK II) 活化,使得原本藉著synapsin I而束縛在細胞骨架上的突觸小泡( synaptic vesicle )因synapsin I被CaMK II磷酸化而釋放出來,增加了突觸小泡被釋放的機率。實驗結果發現在CaMK II抑制劑的處理下,IGF-I的作用明顯被抑制,顯示IGF-I確實能提高細胞內Ca2+而活化CaMK II來增加神經傳導物質釋放的頻率。
綜合以上結果,我們認為IGF-I能夠藉由PI 3-kinase及PLCg路徑的活化而打開IP3及ryanodine sensitive的鈣離子儲存池,使神經末梢內鈣離子濃度升高後,鈣離子經由活化CaMK II pathway而增加神經傳遞物質的釋放。


Successful synaptic transmission at the neuromuscular junction depends on the precise alignment of the nerve terminals with the postsynaptic specialization of the muscle fiber. It is increasingly apparent that this precision is achieved during development and maintained in the adult through signals exchanged between motoneurons and their target muscle fibers that serve to coordinate their spatial and temporal differentiation. Several aspects of neuronal differentiation appear to be dependent on retrograde signals from the target and studies about synaptic modulation have now focused attention on the characterization of proteins that mediate retrograde signals regulating the organization and function of nerve terminals. According to the published evidences, we find Insulin-like growth factor-I (IGF-I ) might be one of these potential factors.
The acute application of IGF-I, a factor which has been addressed to widely express in developing myocyte, dose-dependently enhances the spontaneous acetylcholine secretion at developing neuromuscular synapses in Xenopus cell culture using whole-cell patch clamp recording. The IGF-I-induced potentiating effect is not abolished when calcium is eliminated from culture medium or bath application of pharmacological calcium channel blocker cadmium, indicating calcium influx through voltage-activated calcium channels are not required. We further define the roles of intracellular Ca2+ stores in IGF-I-induced synaptic potentiation. To approach this problem, Ca2+-ATPase inhibitor thapsigargin were initially used to deplete internal Ca2+ stores. IGF-I no longer elicited any changes in SSC frequency in thapsigargin-treated synapses suggesting that an increase in [Ca2+]i due to Ca2+ release from intracellular Ca2+ stores may contribute to the facilitation of transmitter release induced by IGF-I. Application of membrane-permeable inhibitors of IP3-induced Ca2+ release 2-aminoethoxydiphenyl borate (2-APB) or Xestospongin C (XeC) effectively occluded the increase of SSC frequency elicited by IGF-I. Furthermore, pretreatment of the cultures with ryanodine receptor antagonist 8-(dethylamino) octyl 3, 4, 5-trimethoxybenzoate (TMB-8) also blocked the IGF-I effects indicating that IGF-I activates IP3 and/or ryanodine pathway to initiate calcium release from intracellular stores which subsequently potentiate transmitter release. Treating cells with inhibitors of phosphoinositide-3 kinase (wortmannin and LY294002) and Phospholipase C-g (U73122), but not inhibitor of MAP kinase (PD98059) abolishes IGF-1-induced potentiation of synaptic transmission. Inhibition of Ca2+/calmodulin-dependent protein kinase II (CaMKII) by KN-62 effectively blocks the effect of IGF-I. Taken collectively, our results obtained suggest that IGF-I potentiates neurotransmitter secretion by stimulating Ca2+ release from IP3 and ryanodine sensitive intracellular calcium stores via activate PI3 and/or PLC-g signaling cascades, which leading to an activation of CaMKII-dependent transmitter release.


目 錄
             頁數
中文摘要…………………………………………………………………...1
英文摘要…………………………………………………………………...4
緒論………………………………………………………………………...6
Insulin-like growth factor system .................................................................7
IGF-1 and Type 1 IGF receptor ………………………………...............….8
Insulin-like growth factor binding proteins ( IGFBPs )………..................11
IGFs在神經系統的分佈及表現情形…………………………………….12
神經傳導物質的分泌………………………………………….................13
實驗目的……………………………………………………….................16
實驗材料……………………………………………………….................17
實驗方法……………………………………………………….................19
1.電生理記錄方法……………………………………………………....................19
2.實驗數據分析及統計………………………………………………....................20
3.實驗用試劑及供應者………………………………………………....................20
實驗結果………………………………………………………………….21
討論……………………………………………………………………….29
參考文獻………………………………………………………………….33
圖檔順序 頁數
附圖 1…………………………………………………………………….44
附圖 2…………………………………………………………………….46
附圖 3…………………………………………………………………….47
附圖 4…………………………………………………………………….48
附圖 5…………………………………………………………………….49
Fig. 1………………………………………………………………………50
Fig. 2………………………………………………………………………52
Table 1…………………………………………………………………….53
Fig. 3………………………………………………………………………54
Fig. 4………………………………………………………………………56
Fig. 5………………………………………………………………………58
Fig. 6………………………………………………………………………60
Fig. 7………………………………………………………………………61
Fig. 8 .……………………………………………………………………..63
Fig. 9………………………………………………………………………65


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