臺灣博碩士論文加值系統

海洋刺胞動物與胞內共生之渦鞭毛藻建立出豐腴和多樣性的珊瑚礁生態系。然而，我們對於此共生關係及參與其中的分子機制卻不是很清楚。文獻指出，哺乳類HRS (hepatocyte growth factor-regulated tyrosine kinase substrate) 參與了細胞內蛋白質運輸的調控，並且可能參與了自噬小體成熟化 (autophagosome maturation) 的過程。本研究因此希望探討 HRS蛋白在海洋刺胞動物與渦鞭毛藻胞內共生參與的情形。免疫螢光染色分析的結果顯示，大多數美麗海葵宿主細胞內的共生小體含有ApHRS免疫螢光訊號。以DCMU ( 3- ( 3,4-dichlorophenyl ) -1,1-dimethylurea ) 光合作用抑制劑處理海葵，導致ApHRS + 共生小體的比例大幅減少。吞噬作用 (phagocytosis test) 結果顯示，包著酵母菌的胞噬小體 (phagosome) 上帶有ApHRS免疫螢光訊號，而餵食120分鐘之後的胞噬小體，此比例高於餵食60分鐘之後的。進一步選殖了美麗海葵同源性蛋白LC3 (microtubule-associated protein 1 light chain 3，自噬小體標記蛋白)。在胺基酸序列上，ApLC3與人類LC3有72 %的相似性。二重免疫染色實驗發現，ApHRS和ApLC3常同時座落在相同的共生小體微膜區 (symbiosome membrane microdomains) 上。根據以上實驗結果，ApHRS和ApLC3很可能參與美麗海葵和共生藻所建立胞內共生的調控。

The intracellular association of symbiotic dinoflagellates (zooxanthellae) with marine cnidarians is the foundation of the highly productive and diversified coral reef ecosystems. However, the responsible molecular mechanism for its establishment and maintenance is still poorly known. Literature shows that HRS regulates intracellular protein sorting and may play an important role in autophagosomal maturation. The current study aimed to investigate the role of HRS protein in the endosymbiosis between the sea anemone Aiptasia pulchella and zooxanthellae. Immunostaining analysis showed that ApHRS immunoreactivities were associated with the majority of symbiosomes. DCMU treatment of the sea anemone resulted in great reduction in the percentage of ApHRS + symbiosomes within 30 min. However, this percentage rose again when DCMU treatment was prolonged to 120 min. Phagocytosis assay showed that phagosomes containing yeasts were highly decorated with ApHRS-positive immunosignals, with higher preference toward the 120 minutes-old phagosomes than 60 minutes-old phagosomes. A partial cDNA for ApLC3 protein, the sea anemone homologue of LC3 proteins, which is 72 % identical to human LC3 protein, was cloned and employed as an autophagosome marker. Double immunostaining showed that ApLC3 was located to ApHRS-positive symbiosome membrane. Overall, the present study supported that ApHRS is involved in the regulation of Aiptaisia-microalgal endosymbiosis.

目錄.......................................................I
圖目錄............. .......................................IV
附錄目錄...................................................VI
中文摘要....................................................1
英文摘要....................................................3
第一章、 前言...............................................5
1.1光合胞內共生的定義…........................................5
1.1.1 光合胞內共生普遍發生於海洋生物中...........................5
1.1.2 光合胞內共生對珊瑚礁生態系的重要性.........................6
1.1.3光合胞內共生對珊瑚宿主的利益...............................6
1.1.4 光合胞內共生對共生藻的利益...............................6
1.2 光合胞內共生的建立........................................7
1.2.1 胞內微生物的生存機制....................................8
1.2.2 光合胞內共生的維持......................................9
1.2.3 光合胞內共生之共生藻的生長與增殖..........................11
1.3 光合胞內共生崩解的因素....................................11
1.3.1 光合胞內共生崩解的機制..................................12
1.4 光合胞內共生之共生藻生存機制...............................13
1.4.1 胞噬小體和共生小體之間的差異.............................14
1.5 海洋刺胞動物降解共生藻的過程...............................15
1.5.1 走往胞噬小體的降解機制..................................15
1.5.2走往自噬小體的降解機制...................................16
1.6 自體吞噬...............................................17
1.6.1自體吞噬中參與的蛋白質和機制..............................17
1.7 HRS在自體吞噬中的角色....................................18
實驗目的...................................................20
第二章、 材料與方法.........................................21
2.1 動物培養.............................................. 21
2.2 ApLC3 cDNA部分片段的選殖與序列分析........................21
2.3 ApLC3 重組蛋白的建構....................................23
2.3.1 ApLC3 重組蛋白的表現與純化.............................25
2.4 蛋白質定量............................................ 26
2.4.1十二烷基硫酸鈉聚丙烯醯胺凝膠電泳..........................27
2.4.2西方點墨法............................................29
2.5免疫螢光染色.............................................31
2.5.1 玻片製備.............................................31
2.5.2 酸固定液的製備........................................32
2.6 DCMU處理作用...........................................32
2.7 吞噬作用...............................................32
2.8 移除酵母菌反應性抗體之酵母菌餵食實驗........................33
2.9 二重染色...............................................33
2.10 以biotin標定海葵內胚層細胞表面...........................34
第三章、結果...............................................36
3.1 Anti-ApHRS抗體的專一性測試(Western blot analysis)........36
3.1.1海葵細胞的免疫螢光染色(共生小體染上的百分比)................36
3.1.2海葵細胞的免疫螢光染色(DCMU處理的結果)....................37
3.2 ApHRS的二重染色分析.....................................38
3.2.1以biotin標定海葵內胚層細胞表面的實驗分析..................39
3.2.2 ApHRS出現在不同時期的yeast-containing phagosomes的比例.......................................................40
3.3 ApLC3的分子選殖.......................................42
3.4 ApLC3 重組蛋白的西方點墨法..............................43
3.5 ApHRS 與ApLC3的共分佈分析..............................44
第四章、討論...............................................45
4.1美麗海葵ApHRS在海葵細胞分佈的情形..........................45
4.2 DCMU藥劑破壞美麗海葵與共生藻的共生關係......................46
4.3 ApHRS蛋白與胞噬作用的關係................................46
4.4 ApHRS的免疫訊號與biotin-labeled endocytic vesicles關係.......................................................47
4.5使用二重免疫螢光染色來得知ApHRS可能扮演的角色................48
4.6 總結.................................................49
第五章、圖表...............................................50
第六章、參考文獻............................................74
附錄......................................................88
圖1. ApHRS的部分cDNA序列和預測的胺基酸序列........................................................50
圖2. 藉由西方點墨法來分析ApHRS於美麗海葵蛋白質樣品的表現情形........................................................52
圖3. 美麗海葵HRS蛋白(ApHRS)胞內分佈的免疫螢光染色分析........................................................53
圖4. DCMU處理與共生小體染上ApHRS免疫螢光訊號的比例分析........................................................54
圖5.以二重免疫螢光染色法分析美麗海葵宿主細胞內的ApHRS和ApRab5分佈情形........................................................55
圖6. 藉由二重免疫螢光染色法來分析美麗海葵宿主細胞內的ApHRS和ApRab7分佈情形......................................................56
圖7. 藉由二重免疫螢光染色法來分析美麗海葵宿主細胞內的ApHRS和Ub-proteins分佈情形...........................................57
圖8. 藉由二重免疫螢光染色法來分析美麗海葵宿主細胞內的ApHRS和ApRab3分佈情形......................................................58
圖9. 藉由cell surface-bound biotin螢光染色法來分析美麗海葵宿主細胞內的biotin-avidin所標定蛋白質和ApRab5分佈情形.....................................................60
圖10. 藉由cell surface-bound biotin螢光染色法來分析美麗海葵宿主細胞內的biotin-avidin所標定蛋白質和ApRab4分佈情形........................................................62
圖11. 藉由cell surface-bound biotin螢光染色法來分析美麗海葵宿主細胞內的biotin-avidin所標定蛋白質和ApHRS分佈情形.......................................................63
圖12. 藉由酵母菌餵食實驗之免疫螢光染色法來觀察美麗海葵細胞內生性ApHRS分佈情形...................................................65
圖13. 利用酵母菌餵食來執行吞噬作用經由免疫螢光染色法來觀察美麗海葵細胞內生性的ApHRS，在消化細胞內包著酵母菌的胞噬小體膜上分佈趨勢圖表.......................................................67
圖14. 從美麗海葵cDNA library選殖出來的部分LC3序列之PCR產物.......................................................68
圖15. 美麗海葵與其他物種的LC3蛋白之多重序列比對親源分析圖.......................................................69
圖16. ApLC3 樹狀親源關係圖......................................................70
圖17. 藉由西方點墨法來分析ApLC3於美麗海葵蛋白質樣品的表現情形.......................................................71
圖18. 以二重免疫螢光染色法分析美麗海葵宿主細胞內的ApHRS和ApLC3分佈情形.......................................................72
附錄一、藥劑之廠牌和貨號.....................................88
附錄二、pCR-TOPO 2.1 載體示意圖..............................90
附錄三、pET100/D-TOPO載體示意圖..............................91
附錄四、Rab蛋白於細胞內的路徑圖................................92

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