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研究生:鄭倢安
研究生(外文):Cheng, Chieh-An
論文名稱:氨暴露導致斑馬魚胚胎離子調節損傷及成魚行為改變
論文名稱(外文):Ammonia exposure impairs ion regulation in zebrafish embryos and changes behaviors in adult zebrafish
指導教授:林豊益
指導教授(外文):Lin, Li-Yih
學位類別:碩士
校院名稱:國立臺灣師範大學
系所名稱:生命科學系
學門:生命科學學門
學類:生物學類
論文出版年:2021
畢業學年度:109
語文別:中文
論文頁數:76
中文關鍵詞:斑馬魚離子細胞表皮角質細胞細胞凋亡氧化壓力粒線體損傷行為改變
外文關鍵詞:ammoniaapoptosisionocytekeratinocytemitochondria damageoxidative stresszebrafishbehavioral alteration
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氨(包含氣態的NH3以及離子態的NH4+)為魚類代謝胺基酸後產生的主要含氮廢物,也是常見的環境汙染物。當魚體內氨濃度提高,將會導致魚隻中樞神經受損,抽搐、昏迷甚至死亡。然而,目前研究中多著重在高氨處理後魚類的適應機制,關於氨對魚隻離子調節功能及行為的毒性作用尚不清楚。本研究分為兩個部分,首先利用斑馬魚胚胎作為模式動物,探討氨如何對胚胎離子調節功能造成損傷,接著利用斑馬魚成魚作為模式動物,評估氨處理後斑馬魚的行為改變。在胚胎毒性研究中,浸泡於不同濃度(0、10、15、20 mM)的氯化銨溶液中96小時(4-100 hpf)後,觀察胚胎卵黃囊上離子細胞及表皮角質細胞。結果指出,20 mM氨處理後離子細胞內氧化壓力上升(CellROX螢光亮度顯著上升)且由Rhodamine 123標定的具粒線體活性離子細胞數目顯著下降,顯示粒線體活性降低。此外,以細胞免疫螢光染色標定20 mM氨處理後凋亡細胞數目顯著上升,並觀察到表皮角質細胞結構損傷。綜合以上結果發現,在高氨處理下,斑馬魚胚胎離子細胞及表皮角質細胞損傷,導致斑馬魚胚胎失去體表屏障,體內離子大量流失。而在行為實驗中,將斑馬魚浸泡於不同濃度(0、1、5、10 mM)的氯化銨溶液中4小時後,對游泳行為、社交行為、學習與記憶能力等面向進行不同實驗。結果顯示1 mM氨處理時可以促進學習記憶能力;5 mM時焦慮及恐懼程度提升且群游下降;10 mM氨處理時活動力、社交行為及焦慮程度下降,但恐懼程度上升。綜上所述,在不同濃度氨暴露以及不同的環境刺激下,斑馬魚的游泳、社交、學習等行為改變,而這些改變可能使斑馬魚存活率下降,進一步使個體適存度降低。
Ammonia (including NH3 and NH4+) is a toxic nitrogenous product of fish, and also a common environmental pollutant. Accumulation of ammonia is toxic to fishes causing convulsions, coma and death. However, the toxic effects of ammonia on fish ion regulation and behaviors are not fully understood. The 1st purpose of this study was to investigate how ammonia impairs ion regulation in fish embryos, and 2nd purpose was to investigate how ammonia influences behaviors in adult fish. Zebrafish were used in this study because it is a popular animal model for toxicological studies. In embryonic experiments, zebrafish embryos were exposed to NH4Cl (0, 10, 15, or 20 mM) for 96 h (4-100 hpf) and the ionocytes and keratinocytes in the skin were examined. Results showed that 20 mM NH4Cl exposure increased the oxidative stress (indicated by CellROX staining) and decreased mitochondria activity (indicated by rhodamine-123 staining) in the skin ionocytes. Also, 20 mM NH4Cl increased apoptosis (indicated by caspase immunostaining) and impaired the apical structure of keratinocytes. Taken together, high concentration ammonia impaired the skin ionocytes and keratinocytes, causing severe ion losses in zebrafish embryos. In behavior experiments, adult zebrafish were exposed to NH4Cl (0, 1, 5 or 10 mM) for 4 h, and several behaviors including swimming behavior, social behavior, learning and memory were analyzed. The results showed that 1 mM NH4Cl promoted learning behavior; 5 mM NH4Cl increased the anxiety level and fear response, but decreased the shoaling behavior; 10 mM NH4Cl decreased swimming activity, social behaviors, and anxiety level but increased fear response. Taken together, ammonia exposure altered the social behaviors, learning and memory in adult zebrafish.
摘要 1
Abstract 2
研究背景 4
淡水魚氨的生成及排氨機制 4
氨的毒性機制 5
氨對魚類的毒性研究 6
斑馬魚模式動物 7
斑馬魚胚胎表皮角質細胞(Keratinocyte)及離子細胞(Ionocyte) 7
斑馬魚行為實驗 8
氨對魚類行為的影響 9
研究目的 11
實驗流程圖 12
一、胚胎毒性實驗流程圖 12
二、成魚行為實驗流程圖 12
實驗設計 13
實驗1:96小時氯化銨(15 mM)處理後對胚胎基因表現量的影響。 13
實驗2:96小時氯化銨(10、15、20 mM)處理後對胚胎卵黃囊上離子細胞的影響。 13
實驗3:96小時氯化銨(10、15、20 mM)處理對胚胎卵黃囊上具粒線體活性細胞數目的影響。 13
實驗4:72小時氯化銨(15、20 mM)處理後對胚胎卵黃囊上細胞氧化壓力的影響。 14
實驗5:96小時氯化銨(15、20 mM)處理後對胚胎卵黃囊上細胞凋亡的影響。 14
實驗6:96小時氯化銨(15、20 mM)處理後所導致胚胎卵黃囊上表皮細胞凋亡現象。 14
實驗7:96小時氯化銨(15、20 mM)處理後對胚胎卵黃囊上表皮型態的影響。 15
實驗8:96小時氯化銨(5、10、15、20 mM)處理後胚胎逃跑反應。 15
實驗9:4小時氯化銨(1、5、10 mM)處理後對成魚游泳行為的影響。 15
實驗10:4小時氯化銨(5、10 mM)處理後對成魚焦慮程度的影響。 15
實驗11:4小時氯化銨(5、10 mM)處理後對成魚社交行為的影響。 16
實驗12:4小時氯化銨(5、10 mM)處理後對成魚社交認知影響。 16
實驗13:4小時氯化銨(5、10 mM)處理後對成魚膽量的影響。 16
實驗14:4小時氯化銨(5、10 mM)處理後對成魚學習和記憶的影響。 16
實驗15:4小時氯化銨(5、10 mM)處理後對成魚恐懼反應的影響。 17
實驗16:4小時氯化銨(1、5、10 mM)處理後對成魚腦內基因表現量的影響。 17
材料與方法 18
實驗動物 18
氯化銨的配置與處理 18
定量即時聚合酶連鎖反應(Quantitative real time polymerase chain reaction) 19
免疫組織化學染色(immunohistochemistry, IHC) 19
活體螢光染劑 20
離子細胞活體染色標記 21
活性氧化物測定 21
細胞凋亡測定 22
影像分析 22
掃描式電子顯微鏡(Scanning electron microscopy, SEM) 22
觸碰誘發反應(Touch-evoked response assay) 23
新魚缸探索測試(Novel tank diving test) 23
社交偏好測試(Social preference test) 24
社交認知測試(Social recognition test) 24
群游測試(Shoaling test) 25
主動迴避測試(Active Avoidance test) 25
開放空間測試(Open field test) 26
新事物探索測試(Novel object approach test) 26
暗示條件恐懼制約測試(Cued fear conditioning test) 26
統計方法 27
結果 29
實驗1:96小時氯化銨(15 mM)處理後對胚胎基因表現量的影響。 29
實驗2:96小時氯化銨(10、15、20 mM)處理後對胚胎卵黃囊上離子細胞的影響。 29
實驗3:96小時氯化銨(10、15、20 mM)處理對胚胎卵黃囊上具粒線體活性細胞數目的影響。 29
實驗4:72小時氯化銨(15、20 mM)處理後對胚胎卵黃囊上細胞氧化壓力的影響。 30
實驗5:96小時氯化銨(15、20 mM)處理後對胚胎卵黃囊上細胞凋亡的影響。 30
實驗6:96小時氯化銨(15、20 mM)處理後胚胎卵黃囊上表皮細胞凋亡現象。 30
實驗7:96小時氯化銨(15、20 mM)處理後對胚胎卵黃囊上表皮型態的影響。 31
實驗8:96小時氯化銨(5、10、15、20 mM)處理後胚胎逃跑反應。 31
實驗9:4小時氯化銨(1、5、10 mM)處理後對成魚游泳行為的影響。 32
實驗10:4小時氯化銨(5、10 mM)處理後對成魚焦慮程度的影響。 32
實驗11:4小時氯化銨(5、10 mM)處理後對成魚社交行為的影響。 32
實驗12:4小時氯化銨(5、10 mM)處理後對成魚社交認知影響。 33
實驗13:4小時氯化銨(5、10 mM)處理後對成魚膽量的影響。 33
實驗14:4小時氯化銨(5、10 mM)處理後對成魚學習和記憶的影響。 33
實驗15:4小時氯化銨(5、10 mM)處理後對成魚恐懼反應的影響。 34
實驗16:4小時氯化銨(1、5、10 mM)處理後對成魚腦內基因表現量的影響。 34
討論 35
氨處理對斑馬魚胚胎離子細胞及離子調節影響 35
斑馬魚胚胎離子細胞粒線體損傷及過量氧化壓力之關係 35
斑馬魚胚胎表皮角質細胞凋亡與胚胎死亡機制 36
氨處理對斑馬魚胚胎生理功能影響 36
氨處理對斑馬魚成魚活動力及焦慮程度的影響 37
氨處理對斑馬魚成魚社交行為的影響 38
氨處理對斑馬魚成魚學習與記憶能力的影響 39
氨處理後對斑馬魚整體行為影響 40
氨處理斑馬魚作為其他研究模型 41
結論 43
參考文獻 44
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