蛇毒心臟毒蛋白(cardiotoxin)為一系列結構相似的beta-板狀蛋白(beta-sheet proteins)，他們使被咬傷者的心肌收縮而停止跳動，並且引發細胞壞死等現象，但是詳細機制還不清楚。在本論文中，我們提出蛇毒心臟毒蛋白A3 在H9C2細胞株的作用中，細胞膜上的硫脂(sulfatide)聚集區扮演重要角色。蛇毒心臟毒蛋白A3與硫脂結合後不但在細胞膜上產生可導通離子的孔洞，也幫助蛇毒心臟毒蛋白A3進入細胞進而在粒線體聚集。硫脂能夠與蛇毒心臟毒蛋白A3結合並且幫助蛇毒心臟毒蛋白A3穿入細胞膜，促使蛇毒心臟毒蛋白A3在細胞膜上形成不具離子選擇性的孔洞。而蛇毒心臟毒蛋白A3引起之神經醯胺(ceramide)之介面區(interfacial region)脂質結構變化為導致蛇毒心臟毒蛋白A3多體化(oligomerization)與引發毒性的關鍵步驟。硫脂促使的蛇毒心臟毒蛋白A3進入細胞機制為對膽固醇敏感、與溫度相關，但是與三磷酸腺苷、胞膜窖以及籠形蛋白無關的胞飲作用。根據外加的不同鏈長肝素對蛇毒心臟毒蛋白A3進入細胞的促進作用比較，我們提出蛇毒心臟毒蛋白A3雙體化(dimerization)為進入細胞的重要中間產物(intermediate)。蛇毒心臟毒蛋白A3進入細胞後，約於五分鐘後聚集於粒線體，進而裂解粒線體網路(mitochondria network)。蛇毒心臟毒蛋白A3裂解粒線體網路的作用可以在無胞內鈣離子的狀態下進行，但是當胞內鈣離子受到蛇毒心臟毒蛋白A3刺激而濃度上升時，將對粒線體網路裂解有加成作用。而蛇毒心臟毒蛋白A3即藉由針對細胞膜與粒線體膜的作用，使得H9C2肌母細胞株壞死(necrosis)。

Cobra cardiotoxins (CTXs) are structurally related snake venom beta-sheet polypeptides that cause systolic cardiac arrest and severe tissue necrosis of the bitten victim with unknown mechanism. Here, we show that sulfatide (SGC) lipid domain is involved in CTX A3 action in H9C2 cell not only to promote CTX pore formation in plasma membrane but also to facilitate CTX internalization for targeting mitochondria. SGC can interact with CTX A3 to form a non-specific ion conduction pore on cell membrane via a sulfatide-induced insertion of CTX into cell membrane. The CTX-induced lipid conformational change at the ceramide interfacial region is a key step for the action of CTX A3 to induce oligomerization and to cause its cytotoxicity in an SGC-dependent manner. CTX A3 internalization is facilitated by sulfatide in plasma membrane by a cholesterol-sensitive, temperature-dependent, ATP-independent, caveolae-independent, and clathrin-independent, endocytic pathway. Based on exogenous addition of heparin with various chain-lengths, we propose that CTX A3 dimerization is an important intermediate for internalization. CTX A3 targets mitochondria after its internalization in ~5 minutes and disrupts the mitochondrial network in the presence or absence of CTX-induced calcium increase. By action on the plasma and mitochondrial membranes, CTX-induced a necrotic cell death of H9C2 myoblasts.

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