臺灣博碩士論文加值系統

全文摘要
鋅是生物必須之微量元素，一般動物組織中之鋅濃度約為10 ~ 100 μg/g wet weight，然而鯉魚之消化道組織卻有高濃度之鋅 (562 ± 287 μg/g fresh tissue)，遠高於其他魚類。本論文希望了解鯉魚含有高鋅之生理現象與其生態是否有關。鯉魚在生態上與其他生物最不同之處是其能極度容忍缺氧，並能耐極端溫度之變化，鯉魚可以生存在4 ~ 35℃之水溫中。缺氧與水溫劇烈變化對魚類都是一種緊迫 (stress)。本論文係研討鯉魚組織中之高鋅與其能容忍緊迫之相關。
為了解鯉魚及鯽魚之高鋅與能耐缺氧兩者間是否有關，乃進行以下實驗。將五批鯉魚與六批鯽魚 (每批各100尾)，在水溫25 ~ 31℃下，進行缺氧實驗5 d，發現494尾鯉魚平均死亡率為20%，但600尾鯽魚完全沒有死亡。死亡鯉魚消化道組織中之鋅濃度，明顯地比存活之魚低 (39 ± 30 vs 152 ± 44 μg/g fresh tissue, P < 0.001)。鯽魚消化道組織中鋅濃度，平均為 652 ± 458 μg/g fresh tissue。將實驗中一批不耐缺氧之鯉魚餵食高鋅飼料後，進行缺氧實驗，調查高鋅及其活存率之關係。餵食過高鋅飼料 1及2個月之鯉魚，其存活率由0 %增至～50 %，而餵食高鋅飼料6個月之鯉魚，則100 %存活。由此可知鯉魚及鯽魚消化道組織中高鋅與其缺氧活存率，有密切關係。
為了解在緊迫下，魚類血液plasma cortisol及其鋅濃度是否有關，乃比較實驗室靜養鯉魚 (resting fish) 與市售鯉魚 (stressed fish)，其血液plasma cortisol、鋅濃度、及細胞數目之含量。實驗後發現市售鯉魚血液之平均plasma cortisol、血液鋅濃度、細胞數目皆明顯地高於實驗室之鯉魚 (1,522 vs 200 ng/ml；30.1 vs 3.71 μg/ml whole blood；1.76 vs 1.49 ×109 cells/ml whole blood)。且鯉魚血液plasma cortisol與其總鋅濃度之間，有正相關存在。鯉魚頭腎組織之鋅濃度及細胞數，stressed fish顯著地高於resting fish (665 vs 409 μg/g fresh tissue；1.62 vs 0.81 ×109 cells/g fresh tissue)；在腎臟及脾臟間，stressed fish與resting fish，則無顯著差異。在草魚、鰱魚及吳郭魚三種魚類中，市售魚 (stressed fish) 血液平均plasma cortisol濃度亦皆高於實驗室之魚 (resting fish)。然而，此三種魚類之頭腎、腎臟及脾臟鋅濃度，皆在12 ~ 25 μg/g fresh tissue之間。在stressed fish與resting fish之間，鋅濃度沒有差異。
將鯉魚注射cortisol後，plasma cortisol由平均178升為540 ng/ml；血液細胞濃度顯著地上升，由平均1.06升為1.33×109 cells/ml whole blood；總鋅濃度由平均3.91上升至24.0 μg/ml whole blood。鯉魚注射cortisol後，其消化道組織及肌肉之鋅濃度會下降；血液、頭腎、腎臟及脾臟之鋅濃度會上升。肝胰臟、其他內臟組織及骨骼組織之鋅濃度則沒有顯著變化。鋅主要是由消化道組織，少部分由肌肉移往造血組織 (頭腎、腎臟及脾臟) 及血液。將草魚、鰱魚及吳郭魚注射cortisol後，除了plasma cortisol升高外，對此三種魚類之血液及造血組織鋅濃度並未產生影響。
將100尾鯉魚進行5 d的缺氧實驗後發現，鯉魚血液plasma cortisol濃度，在缺氧1 d後，顯著地由150上升至590 ng/ml (接近四倍值)。同時，消化道組織之鋅濃度由216顯著地下降至69 μg/g fresh tissue；頭腎之鋅濃度，由d 0之328顯著地增加至d 3之1,871 μg/g fresh tissue；而脾臟之鋅濃度，是由d 3顯著地增加至d 7。在缺氧之d 5，鯉魚頭腎鋅濃度增加之時，血液細胞含量也顯著地增加。然而當缺氧緊迫解除時，鯉魚之血液、消化道組織、頭腎及脾臟組織之鋅含量則恢復至未缺氧之狀態。在靜養時，鯉魚會將鋅儲存在消化道組織，當遇到緊迫時，鋅會解離，輸送至頭腎；在頭腎中，誘導紅血球增生。增生之紅血球可能是經脾臟而輸送至血液。
將靜養時鯉魚血液製成抹片，經Giemsa染色後，由顯微鏡觀察，可知鯉魚血液之組成主要為紅血球、6 μm細胞及白血球，其所佔比例分別為89%、6%及5%。紅血球及6 μm細胞對43 kDa鋅結合蛋白質之抗體皆有反應，但是白血球則無反應。為了確定6 μm細胞為何種細胞，將鯉魚先行大量抽血 (占其總血量約40 %)，經不同日期後，再抽血觀察其血液之生成。鯉魚在抽血6 d後，血液細胞中6 μm細胞含量由6 %上升至23 %。此增生之6 μm細胞應為新生紅血球。免疫螢光實驗顯示這些6 μm細胞對43 kDa鋅結合蛋白質抗體會呈現反應，且其反應比紅血球強烈。鯉魚在靜養狀態、市售、注射cortisol後及缺氧後，其新生紅血球比例分別為6、48、20、18 %；plasma cortisol濃度則分別為200、1,522、540、460 ng/ml。新生紅血球之比例與其plasma cortisol濃度成正比。餵食高鋅鯉魚血液細胞中，有較一般鯉魚多出數倍之新生紅血球。本論文所研究之市售鯉魚、注射cortisol、遭遇缺氧緊迫及高鋅鯉魚之血液細胞中，都顯示有大量新生紅血球細胞。鯉魚為什麼比其他魚類更容易適應緊迫之主要原因可能是，鯉魚具有一個有效率調控醣皮質接收器之系統。在此系統中，鋅經由醣皮質接收器扮演一個重要之角色。

Summary
Zinc is an essential nutrient for most organisms. The content of zinc in most tissues of different species is on the order of 10 ~ 100 μg/g wet weight. However, the common carp Cyprinus carpio has high zinc contents in its digestive tract tissue (562 ± 287 μg/g fresh tissue) which is not seen in other fishes. The common carp is able to survive days of anoxia at room temperature and can survive for several months at temperatures between 4 ~ 35℃. In this report, the link between high zinc levels and the extreme stress tolerance of common carp was studied.
Five lots of common carp and six lots of crucian carp (each lot of 100 fish) were treated under anoxia for 5 days at a water temperature of 25 ~ 31℃. The average mortality for the whole sample totaling 494 common carp was 20 %. None of the 600 crucian carp in all six groups subjected to 5 days of anoxia died. The zinc concentration in the whole tissue of the digestive tract of the common carp that died was significantly lower than the concentrations found in the sample of fish that survived (39 ± 30 vs 152 ± 44 μg/g fresh tissue, P < 0.001). Mean zinc concentration in the digestive tract tissues of crucian carp just before the start of the experiment was 652 ± 458 μg/g fresh tissue. A lot of common carp which had a low tolerance for anoxia were used to investigate the effect of a high zinc diet on survival from anoxia. Feeding these fish a high zinc diet (2,000 mg zinc/kg diet) for 1 or 2 months increased their survival rate by as much as 50%. Feeding them the same high zinc diet for 6 months brought the survival rate to 100 % over the 5 days of anoxia. It was found the high tolerance to anoxia in common carp and crucian carp is related to zinc concentration in the digestive tract tissue of the fish.
Plasma cortisol, zinc and blood cell contents in the laboratory common carp (resting fish) were compared with those from the market (stressed fish). The plasma cortisol concentration, zinc concentration in the whole blood, and the blood cell concentration from the stressed fish were significantly greater than those from resting fish (1,522 vs 200 ng/ml, 30.1 vs 3.71 μg/ml whole blood, 1.76 vs 1.49 × 109cells/ml whole blood). A strong positive correlation between plasma cortisol concentration and blood cell zinc concentration in the common carp was observed. The zinc content and the cell concentration in the whole tissue of stressed common carp head kidney is significantly higher than that from the resting fish (665 vs 409 μg/g fresh tissue, 1.62 vs 0.81 × 109cells/ml whole blood). However, there were no significant differences between the zinc contents in the whole tissues of common carp kidney and spleen from the stressed and resting samples. The plasma cortisol concentrations in the grass carp, silver carp and tilapia from the stressed fish were all much higher than those from the resting fish. However, the difference of zinc concentrations in the whole blood of these fish are rather small. The mean zinc contents in all of the various tissues analyzed from the different grass carp, silver carp and tilapia are between 12 and 25 μg/g fresh tissue. There was little or no difference between the zinc contents from the stressed and the resting grass carp, silver carp and tilapia.
Two days after cortisol injection, the mean plasma cortisol concentration in the common carp increased from 178 to 540 ng/ml, while the mean blood cell concentration significantly increased from 1.06 to 1.33 × 109cells/ml whole blood. After cortisol injection, the mean zinc concentration in common carp whole blood increased from 3.91 to 24.0 μg/ml whole blood. Significant zinc decreases in the digestive tract tissue and muscle were observed. Similarly, significant increases in blood, head kidney, kidney and spleen zinc levels were found. There was no change in the zinc level of the hepatopancreas or any other visceral or skeletal tissue. A significant amount of zinc was released from the digestive tract tissue, and to a lesser extent from the muscle, into the hemopoietic tissues (i.e., head kidney, kidney and spleen) and blood. The same concentration of cortisol was injected into grass carp, silver carp and tilapia. The mean plasma cortisol level significantly changed in grass carp and silver carp, respectively. Cortisol injection had no effect on the zinc content in the blood, head kidney, kidney and spleen of the grass carp, silver carp and tilapia.
Common carp (100 fish) were subjected to anoxia for 5 days, followed by 4 days recovery. The mean plasma cortisol level increased from 150 to 590 ng/ml 1 day after anoxia (approximately fourfold). It is notable that the zinc content in the digestive tract tissue of the common carp greatly decreased from 216 to 69 μg/g fresh tissue following the increase in plasma cortisol. Additionally, the mean zinc content in the whole head kidney tissue greatly increased from 328 to 1,871 μg/g fresh tissue. Zinc content in the spleen of the common carp showed a different pattern, increasing from day 3 to day 5, and continued to increase until day 7 (i.e., 2 days after recovery). Both the zinc and cell concentration in common carp whole blood and head kidney significantly increased after 5 days anoxia. When the plasma cortisol level recovered to a normal state, the zinc content in the blood, the digestive tract tissue, head kidney, and spleen also recovered. Under normal conditions, zinc is stored in digestive tract tissue. When under stress, the zinc content in the digestive tract tissue of the common carp was released. The released zinc was transported to the head kidney. In the head kidney, the released zinc may induce erythropoiesis. Considering the timing of the change of zinc in head kidney and spleen, it is very possible that the zinc increase in the spleen from day 3 to day 7 came from head kidney. Increased blood cell number may via spleen come to blood.
Under normal conditions, common carp blood cells were stained with Giemsa for light microscopic examination, it was included red blood cells, 6 μm cells, white blood cells and other cells, the proportions of which were calculated to be 89, 6 and 5 %, respectively. The blood cells of the common carp under normal conditions were immunofluorescently stained with an antibody against the 43 kDa zinc-binding protein isolated from common carp digestive tract tissue. The cell surface of both mature red blood cells and the 6 μm cells were immunofluorescently stained, but white blood cells were not. To determine the identity of these 6 μm cells, common carp were subjected to bleeding to produce new red blood cells. Common carp blood was sampled 6 days after bleeding and stained, the percentage of 6 μm cells increased from 6 to 23%. Therefore, it is very probable that the 6 μm cells are ‘‘immature red blood cells’’. These putative immature red blood cells were immunofluorescently stained with the antibody against the 43-kDa zinc-binding protein. The immature red blood cells had much stronger immunofluorescence than mature red blood cells. The percentage of immature red blood cells in the whole blood of common carp under normal conditions was 6%; in the market, cortisol-injected and anoxia-stressed animals, the percentage was 48, 20 and 18%, respectively. The percentage of immature red blood cells in the common carp whole blood was correlated with mean plasma cortisol values (in ng/ml): 200 for fish under normal conditions, 1,522 for market fish, 540 for cortisol-injected fish and 460 for anoxia-stressed fish. A strong positive correlation between the percentage of immature red blood cells and the mean plasma cortisol values was observed. Feeding common carp a high zinc diet (2,000 mg zinc/kg diet) for 6 months increased their immature red blood cells more than under normal conditions. In the present study, common carp that were obtained from the market, cortisol-injected, tested 5 days after anoxia, or fed common carp a high zinc diet, all showed higher blood cell numbers. One important reason why common carp can tolerate more stress than other fish may be because they have a system that regulates the glucocorticoid receptor more efficiently than other fish under stress. In this system, high zinc levels may play an important role via glucocorticoid receptor.

目 錄
全文摘要 1
英文摘要 5

第一章 研究背景與目的 12
一、 研究背景 12
1. 動物組織高鋅之生理意義，目前並不清楚 12
2. 鯉魚組織中高濃度之鋅 12
3. 鯉魚組織中高濃度鋅之生理特性 13
4. 鯉魚之「鋅結合蛋白質」與其「結締組織細胞」 15
二、 研究目的 16

第二章 文獻整理 17
1. 鋅之生物化學及生理學之研究 17
2. 魚類之stress與神經內分泌反應之關係 18
3. 缺氧時動物之對應 19
4. 低氧或缺氧時，魚類中間代謝 (intermediary metabolism) 之變化 20
5. 鋅之增加紅血球對氧之親和力 21
6. 低溫對魚類之影響 22
7. 注射catecholamines或cortisol對魚類intermediary metabolism之影響 23
8. 造血組織 23
9. 紅血球生成組織 25
10. 紅血球的分化 25
11. 紅血球生命週期 27
12. 魚類之血液數值 27
13. 魚類之血液量 29
14. Cortisol與鋅之關連性 29

第三章 鯉魚之耐“缺氧”與其消化道組織鋅濃度之關係 31
一、 前言 31
二、 材料與方法 32
1. 實驗用魚 32
2. 缺氧實驗 32
3. 試魚取樣 33
4. 鯉魚消化道組織之次細胞分劃 (subcellular fractionation) 34
5. 以EDTA抽取消化道組織的鋅 35
6. 鋅濃度之測定 35
7. 已餵食「高鋅飼料」鯉魚，在缺氧時之存活率 35
三、實驗結果 37
1. 缺氧時，鯉魚之存活率與其消化道組織中之鋅濃度 37
2. 缺氧時，鯽魚之存活率與其消化道組織中之鋅濃度 37
3. 以次細胞分劃鯉魚與鯽魚消化道組織中鋅濃度 37
4. 缺氧期間，鯉魚消化道組織中鋅濃度之變化 38
5. 缺氧時，餵食「高鋅飼料」鯉魚之存活率 38
四、討論 40
1. 鯉魚適應缺氧與組織鋅濃度之關係 40
2. 適應缺氧與高鋅飼料之關係 40
3. 缺氧時，鯉魚及鯽魚之泳動行為 41
4. 缺氧時，鯉魚消化道組織鋅濃度之下降 42
五、摘要 43

第四章 鯉魚血液及造血組織鋅濃度與cortisol之關係 44
一、 前言 44
二、 材料與方法 46
1. 實驗用魚 46
2. Cortisol注射鯉魚之實驗 46
3. 鯉魚注射cortisol 46
4. 試魚之麻醉、抽血 47
5. 試魚之血液分劃 48
6. Cortisol之測定 49
7. 鯉魚頭腎、腎臟及脾臟組織之細胞分劃及鋅濃度測定 50
8. 鯉魚肝胰臟、內臟其他組織、骨骼、肌肉之細胞分劃及
鋅濃度測定 51
三、實驗結果 52
1. 鯉魚血液plasma cortisol及鋅之含量 52
2. Resting與stressed鯉魚頭腎、腎臟及脾臟，細胞數與
鋅含量 53
3. Resting與stressed草魚、鰱魚及吳郭魚血液plasma cortisol與
鋅含量 53

4. Resting與stressed草魚、鰱魚及吳郭魚，其頭腎、腎臟及脾臟之鋅含量 54
5. 鯉魚注射cortisol後，血液鋅濃度之變化 54
6. 鯉魚注射cortisol後，魚體各組織間鋅濃度之變化 55
7. 草魚、鰱魚及吳郭魚注射cortisol後，血液及造血組織
鋅濃度之變化 55
四、討論 56
1. 市場購買之魚 56
2. 鯉魚血液中之plasma cortisol濃度 56
3. 草魚、鰱魚及吳郭魚血液中之plasma cortisol濃度 58
4. 鯉魚血液中plasma cortisol之上升，會促使其鋅濃度上升，
而其他魚不會 58
5. 注射cortisol後，鯉魚各組織間鋅之移動 59
五、摘要 60

第五章 缺氧之緊迫，對鯉魚血液及造血組織鋅濃度
之影響 62
一、 前言 62
二、 材料與方法 63
1. 缺氧實驗 63
2. 試魚之麻醉、抽血 63
3. 試魚之血液分劃 63
4. Cortisol測定 63
5. 鯉魚頭腎、腎臟及脾臟組織之細胞分劃及鋅濃度測定 63
三、實驗結果 64
1. 缺氧緊迫對鯉魚血液plasma cortisol濃度、血液細胞數及血液各部分鋅濃度之影響 64
2. 缺氧緊迫對鯉魚消化道組織及造血組織鋅濃度之影響 64
四、討論 66
1. 在缺氧緊迫開始時，鯉魚plasma cortisol含量會急速上升，
消化道組織鋅濃度下降 66
2. 在缺氧緊迫d 3至d 5之時，鯉魚頭腎之鋅濃度增加，隨後
血液細胞數增加 66
3. 在缺氧緊迫之時，鯉魚頭腎組織與腎臟及脾臟組織間鋅濃度
之關係 66
4. 恢復供氧後，鯉魚各組織鋅之變化 67
五、摘要 68

第六章 鯉魚血液細胞在緊迫中之變化 69
一、 前言 69
二、 材料與方法 71
1. 注射cortisol實驗 71
2. 缺氧實驗 71
3. 試魚之麻醉、抽血 71
4. 試魚之血液分劃 71
5. Giemsa 染色 71
6. 鯉魚鋅結合蛋白質之免疫螢光染色 72

三、實驗結果 74
1. 在靜養時，鯉魚血液細胞之顯微照相 74
2. 鯉魚之新生紅血球 74
3. 注射cortisol後，鯉魚紅血球之變化 75
4. 缺氧後，鯉魚紅血球之變化 76
5. 不同緊迫下，鯉魚新生紅血球之百分比與plasma cortisol
之關係 76
6. 餵食一般飼料及高鋅飼料鯉魚之血液細胞 76
四、討論 78
1. 鯉魚之新生紅血球 78
2. 鯉魚新生紅血球與高鋅鯉魚之抵抗缺氧緊迫 80
五、摘要 81

第七章 綜合討論 82
1. 緊迫會造成所有魚類plasma cortisol濃度上升，但只有在鯉魚，plasma cortisol會誘導組織鋅濃度改變 82
2. Cortisol會誘導鯉魚各組織間鋅濃度變化 83
3. 鯉魚消化道組織是鋅之儲藏庫 84
4. 鯉魚頭腎是鋅之利用處 86
5. 在緊迫下，鋅誘發鯉魚紅血球生成所扮演之角色 87

附表及附圖 89
參考文獻 117

參考文獻
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